—- ey HARVARD UNIVERSITY & Library of the Museum of Comparative Zoology hg eel a is u os ( ee en Sees ie i) fii, i i Ree Ath, oF tas Nes CN y nanan nv i ea j owe any Mae ie THE CANADIAN FIELD-NATURALIST Volume 91 1977 THE OTTAWA FIELD-NATURALISTS’ CLUB OTTAWA CANADA , Fae f heet PS ARIE NOV 291977 The CANADIAN®*"*~ PIPLD-NATURALISTE Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 91, Number 1! January-March 1977 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Their Excellencies the Governor General and Madame Jules Léger The objectives of this Club shall be to promote the appreciation, preservation, and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and co-operate with organizations engaged in preserving, maintaining, or restoring environments of high quality for living things. Members of Council* President: Ewen C.D. Todd Elisabeth Beaubien Jo Ann Murray Vice President: Roger A. Foxall WJ. ri coe Ney : : : A.W. Duga era yen Recording Becta: A.J. Erskine . Berl mR, Grew Roger Teves Corresponding Secretary: A. J. Erskine Daleaubite Stanley M. Teeple Treasurer: Pamela J. Sims H.N. MacKenzie C.G. van Zyll de Jong Diane McClymont * This Council is in office until the Annual Business Meeting in January 1977. Correspondence: Address to The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada K1Y 4J5 The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club with the assistance of contributions from the National Research Council of Canada and The Canadian National Sportsmen’s Show. Opinions and ideas expressed in this journal are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Editor: Lorraine C. Smith Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy Associate Editors C. D. Bird W. Earl Godfrey George H. La Roi E. L. Bousfield Charles Jonkel David P. Scott Francis R. Cook J. Anthony Keith Stephen M. Smith A. J. Erskine Charles J. Krebs Robert E. Wrigley Copy Editor: Marilyn D. Dadswell Business Manager: W. J. Cody Production Manager: Pauline A. Smith Box 3264, Postal Station C Chairman, Publications Committee: C. G. van Zyll de Jong Ottawa, Canada K1Y 4J5 Subscriptions and Membership Subscription rates for individuals are $7.00 per calendar year. Libraries and other institutions may subscribe at the rate of $15.00 per year (volume). The annual membership fee of $7.00 includes club publications. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada K1Y 4J5. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Back Numbers Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field-Naturalists’ Club, 1879-1886, and The Ottawa Naturalist, 1887-1919, may be purchased from the Business Manager. All material intended for publication should be addressed to the Editor: Dr. Lorraine C. Smith, Department of Biology, Carleton University, Ottawa, Ontario, Canada K1S SB6 Cover: Arctic hares (Lepus arcticus monstrabilis) photographed by Gerald R. Parker on 5 August 1973 at Mokka Fiord, Axel Heiberg Island, Northwest Territories. See article on page 8. The Canadian Field-Naturalist Volume 91, Number | January-March 1977 Distribution and Abundance of Waterfowl Wintering in Southern Quebec AUSTIN REED and ANDRE BOURGET Canadian Wildlife Service, Environment Canada, P.O. Box 10 100, Ste-Foy, Quebec G1V 4H5 Reed, Austin and André Bourget. 1977. Canadian Field-Naturalist 91(1): 1-7. Distribution and abundance of waterfowl wintering in southern Quebec. Abstract. In January and February of 1974, 1975, and 1976, surveys of wintering waterfowl were conducted through most open-water areas of southern Quebec. These surveys indicated the presence of at least 171 000 ducks, mostly diving and sea ducks in the estuary and Gulf of St. Lawrence. Inland freshwater areas, mainly in the Montreal region, supported many Common Goldeneye ( Bucephala clangula), Common Merganser (Mergus merganser), and Black Duck (Anas rubripes). The most abundant ducks on the estuarine portion were Oldsquaw (Clangula hyemalis), Common and Barrow’s ( Bucephala islandica) Goldeneyes, and Black Duck. In the gulf, Common Eider (Somateria mollissima), Oldsquaw, Common and Barrow’s Goldeneyes were abundant. The area of the estuary and gulf is of international importance as a sea- and diving-duck wintering ground. Further study and close surveillance are required owing to the birds’ apparently great vulnerability to oil pollution and habitat change in a very rigorous climate. Résumé. Aux mois de janvier et février 1974, 1975 et 1976, des inventaires de sauvagine ont été effectués dans la partie sud de la province de Québec. Au moins 171 000 canards, représentés principalement par des canards de mer et des canards plongeurs ont séjourné durant l’hiver dans l’estuaire et le golfe du Saint-Laurent. Dans les eaux douces de l’intérieur, surtout dans la région de Montréal, on a recensé bon nombre de Garrots communs (Bucephala clangula), de Bec-scies communs (Mergus merganser) et de Canards noirs (Anas rubripes). Dans lestuaire, les espéces observées en plus grand nombre furent le Canard kakawi (Clangula hyemalis), le Garrot commun, le Garrot de Barrow ( Bucephala islandica) et le Canard noir. Dans le golfe, l’ Eider commun (Somateria mollissima), le Canard kakawi, le Garrot commun et le Garrot de Barrow étaient presents en abondance. Nos données prouvent que les régions de l’estuaire et du golfe sont parmi les quartiers d’hiver les plus importants du monde, surtout en ce qui concerne les canards de mer et certains canards plongeurs. I] serait souhaitable que lon poursuive des études et qu’on exerce une surveillance étroite sur les populations hivernantes de sauvagine a cause de leur grande vulnérabilité a la pollution par l’huile et en raison des changements qui surviennent dans un habitat en climat trés rigoureux. Several species of water birds overwinter in southern Quebec, principally along the St. Lawrence river, estuary, and gulf. Heavy ship traffic during the winter and expanding urban and industrial development along the shores are posing an ever increasing threat to those birds and to their habitats. The paucity of published information prompted us to assemble recent un- published reports and to conduct surveys to document the distribution and abundance of waterfowl inhabiting that area during the winter. Study Area and Methods In early February 1974, 1975, and 1976, we attempted to cover all open-water areas in southern Quebec by ground and/or aerial counts. Regional differences in accessibility, habitat type, and species composition prevented the use of a standard survey procedure for the entire study area. We subdivided the area into 10 zones based on these regional differences (Figure 1). The 1974 survey was carried out from 31 January to 20 February, in 1975 from 3 to 6 February, and in 1976 from | to 12 February. Ground crews (P. Blais, A. Bourget, H. Briard, G. Chapdelaine, P. Dupuis, G. Fortin, P. Lamothe, M. Laverdiere, D. Lehoux, P. Rancourt, A. Reed, L.-G. de Repentigny, J. Rosa, J.-P. Savard, and G. Tremblay) 2. THE CANADIAN FIELD-NATURALIST HED, / Z i; (fe LACHINE RAPIDS <7 QUEBEC é Yi, LAKE ST-LOUIS 1G0_ MILES 60 KM F—— ornteans ! ISLAND ONTARIO FIGURE |. Vol. 91 LEGEND Numbers of ducks, all s SEPT -ILES ae NEWFOUNDLAND Cot fice E ISLANOS PRINCE Cuan ISLA SE 4 Southern Quebec, showing major duck wintering sites (dark circles) and survey zones (outlined and identified by large letters). The Montreal area is shown in a larger scale on the inset at the upper left of the figure. The survey zones are (A) Ottawa River, (B) Montreal, (C) Eastern Townships, (D) Lake St. Peter, (E) St. Lawrence estuary, (F) North shore, Gulf of St. Lawrence, (G) Gaspésie, (H) Baie des Chaleurs, (1) Matapedia, (J) Lake St. John. methodically scanned all open-water areas with telescopes and binoculars in all zones except part of zone F. Dupuis and Tremblay conducted aerial surveys that covered mainly zones E and F. In most flights a Cessna 337 was flown at an altitude of approximately 60 m parallel to, and about 70m seaward from, the edge of the shoreline of permanent ice. At some coastal sites flights were conducted up to 15 km from shore to check for offshore flocks of sea ducks. Supplementary observations were available from previous years for the Montreal area and the north shore of the estuary, mainly since 1964. Files of the Canadian Wildlife Service and the Quebec Wildlife Service yielded unpublished results of various winter surveys conducted from 1952 to 1963, mainly in the Montreal area. The winter season was considered to extend from | January to 28 February. Results The combined results of the 1974, 1975, and 1976 surveys are presented in Table | and Figure 1. We believe that we consistently under- estimated the bird populations because (1) exhaustive surveys were not possible over sucha vast and partly inaccessible area, (2) many of the species inhabited offshore areas and were dif- ficult to detect from aircraft (see Stotts and Olson 1972), and (3) cold air temperatures and ice conditions reduced the efficiency of the ob- servers. For these reasons, the largest number of birds recorded in each zone over the three winters can be considered as the best estimate of that population; we have presented those maxi- mum counts in Table 1. The data, in summarized form, are plotted in Figure | to show major concentration areas. Those results, and previous data, form the basis for the description which follows for each area. (A) Ottawa River Most of this area was frozen over in winter; only a few natural pools and areas below a hydroelectric dam remained open. In 1974 (no counts in 1975-76) three species of ducks occurred but only in small groups: Black Ducks (Anas rubripes), Common Goldeneyes (Buce- 1977 REED and BOURGET: WATERFOWL WINTERING IN SOUTHERN QUEBEC TABLE |—Estimated numbers of aquatic birds that overwinter in the St. Lawrence River system, Quebec. Each number represents the maximum count obtained. Surveys were conducted in most regions in February 1974, 1975, and 1976 Regions i) & o ts) = < 3 2 as z oe 65 2 = Species 2 r= ons s = 2 1 ie 8 @ oS Total 2.2 S a0 5 Se aS ge SS gs Ow = oe Ww Za () fea} (OG) 3 642) Black Duck 56 506 71 1906 4 2543 (Anas ribripes) Common Goldeneye 160 2908 164 8328! 979! 332 111 12 8 13002 (Bucephala clangula) Barrow’s Goldeneye l 4 1394 869 260 19 2547 (Bucephala islandica) Bufflehead l 2 69 l 73 (Bucephala albeola) Oldsquaw 6451 2411 46782 48 55692 (Clangula hyemalis) Surf Scoter 185 185 (Melanitta perspicillata) Common Eider 9 91035 505 91549 (Somateria mollissima) Common Merganser 16 5300 196 19 5 5) 7 2 5550 (Mergus merganser) Red-breasted Merganser 2 91 74 98 23 l 289 (Mergus serrator) Miscellaneous | YP 12 3 l 89 duck species? Sub total — Ducks 234 8787 451 18270 95554 47983 206 24 10 171519 Great Cormoran 21 91 112 (Phalacrocorax carbo) Black Guillemot 78 57 42 177 (Cepphus grylle) Total 234 8787 451 18348 95554 48061 339 24 10 171808 'May include some B. islandica. 2Miscellaneous duck species by order of importance are as follows: Ottawa River, Mallard (Anas platvrhynchos), Montreal, Mallard, Pintatl (Anas acura), Canvas- back (A ythya valisineria), Redhead (Avthyva americana), Lesser Scaup (Avthva affinis), Ring-necked Duck (Avhva collaris), American Wigeon (Anas americana). Eastern Townships, Mallard, Hooded Merganser (Lophodvies cucullatus); Estuary, Harlequin Duck ( Histrionicus histrionicus), Mallard: North Shore, Black Scoter (Melanitta nigra). phala clangula), and Common Mergansers (Mergus merganser). (B) Montreal This region had large areas of open water, principally at the inlets and outlets of its two largest bodies of water, Lake St. Francis and Lake St. Louis. The Lachine rapids were partic- ularly important. The Montreal region posses- sed the most important winter-bird concentra- tions in freshwater areas in Quebec. The Common Merganser was the most abundant species in the Montreal region; more than 3500 birds spent the winters of 1974 and 1975 in open water below the Cornwall hydroelectric dam, feeding on fish which had passed through the turbines. The Common Goldeneye was second in importance and the Black Duck third. (C) Eastern Townships All areas south of Lake St. Peter and east of Montreal were included in this zone. 4 THE CANADIAN FIELD-NATURALIST Open-water areas were found along fast-flowing streams and rivers in this rolling countryside. In contrast to the ice-free areas of the St. Law- rence, which were relatively consistent in size and location from year to year, open water was highly variable in extent and location within and between winters. No large concentrations of ducks were encountered. Common Mergansers, Common Goldeneyes, and Black Ducks occur- red regularly. (D) Lake St. Peter This zone had no appreciable areas of permanent open water and no waterfowl were observed. (E) Estuary This portion of the St. Lawrence had large expanses of open water throughout the winter. Of particular importance were large sections of tidal flats near the mouth of the Saguenay River where tides and river currents prevented the formation of permanent ice. Along most of the south shore, water areas suitable for aquatic bird use were frozen over. This vast area included some of the most important wintering sites for waterfowl in the province, especially along the north shore of the St. Lawrence. The most important con- centration of wintering Black Ducks in Quebec was found on the tidal flats near the mouth of the Saguenay River, where more than 75% of the Black Ducks counted in 1974 were found. The Common Goldeneye was the most widely dis- tributed duck on the north shore of the estuary and was the most abundant species. The Old- squaw (Clangula hyemalis) was also important but it was difficult to appraise its distribution and abundance because it often frequented areas too far from shore to be seen by ground crews and because it took flight early and dispersed rapidly at the approach of an aircraft; it was undoubtedly more abundant than our figures indicated. The Barrow’s Goldeneye ( Bucephala islandica) was also encountered frequently. Buffleheads (Bucephala albeola) occurred reg- ularly (although in small numbers) only near the mouth of the Saguenay River. Two other species of aquatic birds also occurred regularly: the Red-breasted Merganser (Mergus serrator) and the Black Guillemot (Cepphus grylle). Vol. 91 (F) North Shore of the Gulf This region extends from Pointe des Monts to Blanc Sablon on the Quebec-— Labrador border (about 800 km) and includes Anticosti Island. Because of the vastness of the area and inaccessibility of certain portions, its coverage cannot be considered exhaustive. The inner part from Pointe des Monts to Matamek was surveyed both from the ground and from the air; east of Matamek was surveyed only from an aircraft. Ice conditions are generally heavy in this region although the area of Natashquan and the eastern tip of Anticosti Island is character- ized by less severe conditions (Simpson 1973). Sea ducks represented the most impor- tant component of the wintering population of aquatic birds. The Common Eider (Soma- teria mollissima) was recorded along the outer north shore of the gulf; most birds of this species were seen in the Jacques-Cartier passage in areas of slushy ice. The Oldsquaw and Goldeneyes (Common and Barrow’s combined) were also abundant. (G) Gaspésie Ice conditions in this zone were highly variable. Water areas were completely covered by large sheets of floating ice when inshore winds prevailed. The eastern tip of the penin- sula, however, was more consistently ice-free; most large flocks were observed there. The Oldsquaw was by far the most abundant bird species observed in 1974 and 1976; most birds of this species were in Baie de Gaspé, which was clear of ice. In 1975, that site was choked with ice and only a small number of Oldsquaws was observed in the entire zone. Similarly, our counts of goldeneye (Common and Barrow’s) varied from year to year, ap- parently reflecting different conditions of ice cover. (H) Baie des Chaleurs This large bay was characterized by heavy ice conditions (Simpson 1973), but oc- casionally strong west winds opened up large expanses of the bay. Despite different ice conditions (1974 heavy, 1975 and 1976 light) few birds were observed. Common and Barrow’s Goldeneyes, Oldsquaw, Red-breasted (Mergus serrator) and Common Mergansers, and Great 1977 Cormorants (Phalacrocorax carbo) were en- countered regularly in all years. (1) Matapedia Many sections of the river remained open through the winter, but were not heavily used by wintering waterfowl. The Common Goldeneye was the main species encountered. (J) Lake St. John The only open-water areas in this zone were found on fast-flowing sections of rivers, principally the Ashuapmuchuan and the Mis- tassini which flow into the lake. Very few birds were present in this area in 1974 (no census in 1975 and 1976). Common Goldeneyes and Common Mergansers were the only species observed. Discussion The abundance and diversity of the wintering population is remarkable for an area withsucha harsh and rigorous climate. In winter waterfowl require open water that can offer an abundant food supply and suitable resting areas (Nilsson 1972). Several species can be accommodated in southern Quebec because there is a wide variety of habitats, ranging from inland-freshwater to coastal-saltwater, which are kept open by tides, winds, and currents. Food is available in the form of aquatic organisms, which are notably abundant in intertidal and sub-littoral zones in the St. Lawrence estuary and gulf. Of the nine species of waterfowl which wintered regularly in this region the Common Eider, the Oldsquaw, and the Common Golden- eye were, in that order, the most numerous. Large flocks of eiders were present in saltwater along the north shore of the gulf, particularly off Natashquan, Baie Johan-Beetz, Havre St- Pierre, and around Anticosti Island. Flocks of several thousand Oldsquaw were found in salt and brackish waters in Baie de Gaspé, the estuary, and along the north shore of the gulf. Common Goldeneyes occurred in fresh, brack- ish, and saltwater habitats, the largest con- centrations being located in the vicinity of the mouth of the Saguenay River (north shore of the estuary) and in the Lachine Rapids near Mont- real. The great abundance of Common Goldeneye REED and BOURGET: WATERFOWL WINTERING IN SOUTHERN QUEBEC 5 and Oldsquaw in the estuary and gulf was heretofore unrecognized (Bellrose 1976; Johns- gard 1975). Similarly, Barrow’s Goldeneye was believed to be relatively rare on the east coast of North America (Bellrose 1976; Johnsgard 1975; Palmer 1976; Hasbrouck 1944): our data in- dicate that the St. Lawrence estuary and gulf represent a stronghold of an unexpectedly large population. Our counts in January and February (1974-1976) indicated an annual wintering pop- ulation of about 171000 ducks, despite in- complete coverage. Clearly our eider estimates must be very low owing to the inaccessibility of many of the known or suspected haunts of this bird; the same applies to the Oldsquaw, with the added complication of its known ability to escape detection by aerial observers (Stotts and Olson 1972). The estimates of goldeneye from coastal areas are undoubtedly low as well. Although precise adjustments cannot be made at this time, it seems likely that the area covered must harbor at least a quarter of a million ducks and it is conceivable that half a million could be involved. Clearly the most important part of the study area was the estuary and gulf of St. Lawrence (Zones E, F, G, and H), which accounted for 94% of the ducks recorded. Some areas of the gulf which were not covered by this study also support important populations of wintering ducks. Average winter populations (1972-1974) on Prince Edward Island were 3070 Black Ducks, 1540 goldeneye (apparently almost all B. clangula), 1450 mergansers (species not indi- cated), and small numbers of Oldsquaw (Bate- man, M. 1974. Mid-winter aerial waterfowl surveys of the Maritime Provinces. Unpublished report, Canadian Wildlife Service, Sackville, N.B. mimeo. 13 pp.). The Magdalen Islands support a few hundred Oldsquaw (A. Smith, personal communication), while Newfoundland is an important gathering area for Common Eiders (Gillespie and Learning 1974). Unfortu- nately there are too many gaps in the data to allow estimation of the winter duck population of the entire gulf. Other areas along the Atlantic coast of North America are also important wintering areas. Various data from aerial surveys in Nova Scotia, 6 THE CANADIAN FIELD-NATURALIST New Brunswick (Bateman, Joc. cit.) and the Atlantic Flyway (winter surveys coordinated by the United States Fish and Wildlife Service) for selected species are presented in Table 2 along with comparative data from the present study. Although those more southerly areas harbor large numbers of surface-feeding ducks it is evident that the St. Lawrence is of great importance to sea and diving ducks. Comparison with European counts is more difficult because a more complete coverage is obtained there by using a network of ground observers (Atkinson-Willes 1969). In terms of total ducks present, however, few areas of equivalent size harbor as many ducks as the St. Lawrence. For example, a comparison of our figures with those from the Baltic Sea, indicated by Atkinson-Willes (1969, Figure 2, p. 105), suggest wintering duck populations of similar magnitude. Also, much of the North Sea has fewer ducks than does the St. Lawrence (Atkinson-Willes 1969; Milne and Campbell 1973). That area of the North Sea encompassing Denmark, the Netherlands and northern Ger- many, however, has a much larger overwinter- ing duck population. The latter area is probably the only European site of greater importance than the St. Lawrence to diving and sea ducks (Atkinson-Willes 1969; Joensen 1974). Clearly the St. Lawrence must be considered one of the major duck wintering areas of the North Atlantic. Voleoi Industrial and urban development along the St. Lawrence pose a constant threat to the birds and to their habitats. Port facilities for super- tankers have been proposed for the estuary and gulf, which would increase the likelihood and gravity of oil spills which could have disastrous effects. In the past two years, 102 cases of oil pollution have been reported in these regions; it is only through good luck that none has had serious effects. In the Lachine Rapids near Montreal, a hydroelectric dam has been pro- posed which, through major changes in the hydrography and biology of the area, could lead to drastic reduction in the Common Goldeneye population. Some habitat changes may lead to an increase in the numbers of some species. The proposed Lachine Rapids dam might increase the numbers of Common Mergansers, as has occurred further upriver near Cornwall. The Lake St. Peter area, which presently has no open water, may eventually serve as a new wintering ground if warm water effluents from nearby nuclear power plants at Gentilly prevent the freezing over of expanses of the St. Lawrence in that area. This investigation has permitted the identi- fication of the more important areas of the system and provided an approximate estimate of numbers of the various species of wintering birds, but further work is required. In partic- ular, the surveys along the remote outer north shore cannot be considered as exhaustive. TABLE 2—Summary of counts of selected species of wintering ducks on the Atlantic coast of North America Nova Scotia and United States} New Brunswick2 Species St. Lawrence! Black Duck 2500 (Anas rubripes) Common Goldeneye 13000 (Bucephala clangula) Oldsquaw 55700 (Clangula hyemalis) Common Eider 91500 (Somateria mollissima) 11800 304500 4400 68200 850 12200 2100 72600 'This study, based on maximum of mid-winter counts 1974-1976 (from Table |, rounded to nearest 100). *Bateman (1974)—Jan. and Feb. 1974 (Atlantic and Fundy coasts). ‘Unpublished results of mid-winter waterfowl surveys in the Atlantic Flyway, coordinated by the United States Fish and Wildlife Service. Average 1964-1973. 1977 Further work is proposed to obtain more accurate counts of sea ducks in the estuary and gulf, to document further the present status of the Barrow’s Goldeneye in winter, and to clarify the racial status of Common Eiders (two races, Somateria mollissima dresseri and S. m. bor- ealis, are. known to overwinter in the gulf (Ouellet 1969, 1975; Gillespie and Learning _1974)). Also it is hoped that a survey can soon be undertaken to cover the entire gulf. The con- tinued and increasing threats from industrial and urban develcpment provide ample justifica- tion for close surveillance and further study. Acknowledgments We are extremely grateful to H. Ouellet, National Museum of Natural Sciences, Ottawa, to M. Lepage, Quebec Wildlife Service, Mont- real, and to Y. Lafleur, Parks Canada, for providing data from unpublished reports. Thanks are due to the many wildlife tech- nicians, pilots, and volunteers who participated in the censuses, often under very difficult con- ditions. The Ministry of Transport kindly permitted us to place an observer on board its helicopter during a lighthouse inspection in the gulf in 1974. H. Ouellet, H. Boyd, and J. Bryant made valuable comments on the manuscript. Literature Cited Atkinson-Willes, G. L. 1969. The mid-winter distribution of wildfowl in Europe, northern Africa and south-west Asia, 1967 and 1968. Wildfowl 20: 98-111. REED and BOURGET: WATERFOWL WINTERING IN SOUTHERN QUEBEC 7 Bellrose, F.C. 1976. Ducks, geese and swans of North America. Stackpole Books, Harrisburg, Pennsylvania. 544 pp. Gillespie, D. I. and W. J. Learning. 1974. Eider numbers and distribution off Newfoundland. /n Canadian Wildlife Service waterfowl studies in eastern Canada 1969-73. Edited by H. Boyd. Canadian Wildlife Service Report Series Number 29. pp. 73-78. Hasbrouck, E. M. 1944. The status of Barrow’s Goldeneye in the eastern United States. Auk 61: 544-554. Joensen, A. H. 1974. Waterfowl populations in Denmark, 1965-1973. Danish Review of Game Biology 9(1). 206 pp. Johnsgard, P. A. 1975. Waterfowl of North America. In- diana University Press, Bloomington, Indiana. 575 pp. Milne, H. and L. H. Campbell. 1973. Wintering sea-ducks off the east coast of Scotland. Bird Study 20: 153-172. Nilsson, L. 1972. Habitat selection, food choice, and feeding habits of diving ducks in coastal waters of South Sweden during the non-breeding season. Ornis Scandi- navica 3: 55-78. Ouellet, H. 1969. Les oiseaux de I’Ile d’Anticosti, province de Québec, Canada. Musée national des sciences naturelles, publications en zoologie Numéro |. 79 pp. Ouellet, H. 1975. Contribution a l’étude des oiseaux d’hiver au Parc National de Forillon, Québec. Revue Géogra- phique de Montréal 29(4): 289-304. Palmer, R. S. (Editor). 1976. Handbook of North Ameri- can birds. Volume 3, Waterfowl (Part 2). Yale University Press, New Haven, Connecticut. 560 pp. Simpson, W. 1973. Gulf of St. Lawrence water uses and related activities. Lands Directorate, Environment Canada, Geographical Paper Number 53. 20 pp. Stotts, R.S. and D.P. Olson. 1972. An evaluation of waterfowl surveys on the New Hampshire coastline. Journal of Wildlife Management 36(2): 468-477. Received 12 May 1975 Accepted 21 November 1976 Morphology, Reproduction, Diet, and Behavior of the Arctic Hare (Lepus arcticus monstrabilis) on Axel Heiberg Island, Northwest Territories GERALD R. PARKER Canadian Wildlife Service, Box 1590, Sackville, New Brunswick EQOA 3C0 Parker, Gerald R. 1977. Morphology, reproduction, diet, and behavior of the arctic hare (Lepus arcticus monstrabilis) on Axel Heiberg Island, Northwest Territories. Canadian Field-Naturalist 91(1): 8-18. Abstract. Fifty-one arctic hares (Lepus arcticus monstrabilis) were collected at Mokka Fiord, Axel Heiberg Island, Northwest Territories: 35 in summer 1973 and 16 in late winter 1975. Adult female weights averaged 4.5 and 3.9 kg for the summer and winter periods respectively; male weights, 4.1 and 4.0 kg. There were significant decreases in the weights of the heart and kidneys between summer and winter samples, as well as an overall loss in body weight. In 1973 the peak of births was believed to have been approximately 20 June. The average litter size, as determined from counts of corpora lutea was approximately five. Breeding occurred about | May; the period of gestation was approximately 50 days. Arctic hares displayed great diversity in their summer diet, but fed mainly on arctic willow (Salix arctica), Dryas integrifolia, and grasses. Arctic willow was the main species consumed at all seasons, and made up 95% by weight of the winter diet. The abundance of willow, a light covering of winter snow, and broken terrain providing adequate escape cover may explain the high densities of arctic hares on parts of Axel Heiberg and Ellesmere Islands. The arctic hare (Lepus arcticus) is found in Canada north of the treeline to the northernmost point of land on Ellesmere Island, Northwest Territories, and also on the rock-strewn plateaus and mountains of eastern Newfoundland. In Greenland it is common on most of the ice-free coastal region. In North America studies of L. arcticus have been limited to short notes on their distribution (Bergerud 1967; J. G. Inder. 1972. Arctic hares on Brunette Island. Typewritten report to New- foundland Wildlife Service. 3 pp.; Watson 1954), taxonomy (Handley 1952; Howell 1936; Nelson 1934), predation (Tener 1954), and natural history (Walkinshaw 1947). Bonnyman (1975) made observations on the behavior and habitat use of hares on the Fosheim Peninsula, Elles- mere Island in the summer of 1975, but detailed information on the biology of the arctic hare is absent from the literature. Arctic hares occur in unusually high densities and often form herds of several hundred or more individuals on parts of Ellesmere and Axel Heiberg Islands. In the summer of 1973 I observed and collected speci- mens of L. arcticus monstrabilis during a study on the feeding habits of muskoxen (Ovibos moschatus) and caribou (Rangifer tarandus pearyi) at Mokka Fiord, Axel Heiberg Island, and returned to Mokka Fiord in late winter 1975 for further observations and collections. Study Area Mokka Fiord is close to the northern limit of arctic hare range (Figure 1). The study area is a valley 6 to 12 km wide; a mountain range is to the west and a lower series of gypsum pierce- ment domes on the east drop abruptly to the waters of Mokka Fiord and Eureka Sound. The valley floor is approximately 200 m above sea- level. The gypsum domes present a region of sparsely vegetated gravel- and rock-strewn ridges and slopes, giving way to an interspersion of gravel ridges and meadows in the valley proper. Meadows are restricted to the glacier-fed streams originating in the mountains to the west. The slopes west of the valley support Salix arctica, Luzula nivalis, Poa spp., Alopecurus alpinus, Arctagrostis latifolia, Dryas integri- folia, Saxifraga spp., and a variety of other forbs. The gravel ridges in the valley support a wide variety of forbs and several grasses; Carex stans, Eriophorum spp., and mosses dominate the meadows. The gravel- and rock-strewn slopes east of the valley support only occasional tufts of grasses (Poa spp., Puccinellia spp.), Carex rupestris, Kobresia myosuroides, and LOTT PARKER: ARCTIC HARE ON AXEL HEIBERG ISLAND 9 300 Miles os Kilometre | FIGURE 1. The location of Mokka Fiord, Axel Heiberg Island, Northwest Territories. scattered mats of Salix arctica. Occasional forbs include Papaver radicatum, Oxyria digyna, Saxifraga oppositifolia, and Dryas integrifolia. Lichens are common but not abundant, the dominant species being Cetraria nivalis, C. cucullata, Thamnolia vermicularis, and Parmelia spp. The mountains of Axel Heiberg Island and eastern Ellesmere Island present a barrier to moist air from the south. This topographical peculiarity contributes to less cloud cover and higher temperatures in summer on western Elles- mere and eastern Axel Heiberg Islands than in most of the Arctic. At Eureka on northwestern Ellesmere Island, mean annual snowfall is 37.5cm and total precipitation only 6.8 cm (Thompson 1967). Mean daily winter tempera- tures are among the coldest in the Canadian Arctic; the average for November to March is —26.9°C. Methods Hares were collected from 3 July to 3 August 1973 and from 23 March to 6 May 1975. They were shot at close range (5 to 20 m) witha .22- caliber rifle. Most specimens were shot in the neck to avoid damage to the skull or internal organs. Each specimen was weighed on spring scales and standard body measurements were taken (Anderson 1965). The heart, lungs, full stomach, liver, and kidneys were weighed on a beam balance. Stomach samples were preserved in an alcohol-formalin-acetic acid solution (AFA) for future analysis. Reproductive tracts were also preserved in AFA. The skulls and mandibles were retained, and a small collection of skins was made from the 1975 collection. All material was deposited with the Canadian Wildlife Service, Atlantic Region, Sackville, New Brunswick. General observations on hare behavior were recorded whenever possible. 10 THE CANADIAN FIELD-NATURALIST In the laboratory, basic cranial and mandib- ular measurements were taken for each speci- men, the specifications conforming with those by Banfield (1961) for caribou. Ovaries were sectioned for a count of corpora lutea. Ovaries were dehydrated and embedded in paraffin, and by use of a rotary microtome, were sectioned in units of 10 uw thick. Every tenth section was mounted on a slide and stained with Weigert’s haematoxylin. External measurements of each ovary were taken in addition to the maximum length, breadth, and depth (number of sections in which structure appears X 10 4) of each corpus luteum. Differences between measurements for com- ponents of the sample were tested for signifi- cance using a simple /-test. The diet of hares was determined by the identi- fication of plant epidermal tissue in stomach samples. The preparation of plant reference slides and key and the method of stomach sample analysis is described by Parker et al. (Parker, G. R., B. Campbell, and M. Gauthier. 1976. Estimating the diet of Arctic herbivores by rumen and faecal analysis. Canadian Wildlife Service Report CWSC-2021. Ottawa. 143 pp.). In contrast to the herbaceous-dominated summer stomach samples, the winter diet con- tained a large amount of unidentifiable woody material. As the only woody plant of any con- sequence is the arctic willow, most of the woody component of the diet was assumed to be that species. Twigs of Dryas integrifolia and the roots of forbs may also have comprised an unknown portion of the non-herbaceous diet. To quantify the importance of herbaceous and Vol. 91 woody forage in the winter diet, a wet sample of 15 to 20 g was washed through a series of sieves of the following dimensions: 1.18 mm; 850 yp; 425 yu; and 300 uw. Samples of material from each sieve were examined for presence of herbaceous material. Only the sieve of 300 u contained appreciable amounts of herbaceous material; the others contained woody fragments almost ex- clusively. Contents of the 300- sieve were then analyzed for plant species composition similar to the summer samples. The weights of material in all sieves were totalled and divided into the weight of herbaceous material in the 300-p sieve. The latter weight was estimated by multiplying the total weight in the 300-u sieve by the percentage of herbaceous fragments determined from microscopic examination. This method is believed to give a close approximation of the importance of woody plant material in the winter diet of arctic hares. Results General Morphology Thirty-five arctic hares were collected at Mokka Fiord, Axel Heiberg Island in July and August 1973, and 16 from March to May 1975. The summer collection consisted of 10 adult males, 15 adult females, 6 juvenile males, and 4 juvenile females. The winter collection was composed of eight males and eight females. I did not distinguish between adults and young-of- the-year in the winter collection. Summer and winter weights and measure- ments of adult hares are shown in Table 1. Inthe summer collection adult females were signifi- cantly heavier (P< 0.01) and of greater girth than adult males. Females collected in the winter TABLE |—The mean weights and measurements, with 95% confidence limits, of adult arctic hares ( Lepus arcticus) collected at Mokka Fiord, Axel Heiberg Island during the periods 3 to 29 July 1973, and 23 March to 6 May 1975 Sex Sample Weight Length Girth Hindfoot Shoulder height Ear size (kg) (mm) (mm) (mm) (mm) (mm) 1973 Female 15 455 +0.7 675.6 + 404 345.6 + 38.4 156.3 + 11.2 3660 +4 42.2 83.6 + 5.4 Male 10 413 +05 669.0 28.8 338.0 + 29.8 154.6 + 8.4 355:5\ = 28:4) 826 e=e5 22 1975 Female 8 3.99 + 0.4 691.2 + 50.1 SWPis) 2 SA) 153.6 + 6.2 388.7 + 52.2 81.9 + 2.4 Male 8 408 + 0.2 690.6 + 38.8 361.9 + 44.1 150.2 + 9.4 386.4 + 35.2 83.0 + 5.7 1977 weighed less than adult females from the summer collection (P < 0.05). Weights for immature hares collected in July and August 1973 and a projected growth curve for the first six weeks of life are shown in Figure 2. Although adult females were generally heavier than adult males, the juvenile weights suggest that males put on weight more rapidly than females during the first six to eight weeks of life. Cranial and mandibular measurements were taken whenever the condition of specimen ma- terial permitted. There was no trend evident in seasonal differences in those measurements. Seasonal weights of selected internal organs of males and females are shown in Table 2. There were significant decreases in mean weights of the heart between summer and winter samples of both adult males (P< 0.05) and adult females (P< 0.05). Both sexes also showed significant decreases in kidney weight from summer to 2.80 (KILOGRAMS) -80 WEIGHT -60 .40 Wt. at birth on June 20 100 grams 28 30 1 4 8 12 20 24 = JUNE ———=] PARKER: ARCTIC HARE ON AXEL HEIBERG ISLAND 11 winter (males, P< 0.05; females, P< 0.001). The liver also showed a seasonal loss of weight although the loss was significant (P < 0.01) only for females. Mean stomach weights of males were significantly (P< 0.001) heavier in the winter than in the summer. In females the mean stomach weight in winter was greater than that of summer but not significantly so. Winter increases could be attributable to a greater intake of woody material. The mean weights of lungs of both sexes were heavier in summer than in winter, although those differences were not significant (P > 0.05). Reproduction The ovaries of 15 adult female arctic hares collected in summer 1973 were sectioned and examined for corpora lutea of pregnancy. All females had ovulated and bred the previous breeding season. The mean number of corpora lutea for the sample was 6.5 (SD = 1.7). The mean ee e MALES * FEMALES 20 24 28 16 iia 4 8 JULY |— AUGUST —4 FIGURE 2. Weights of juvenile arctic hares collected at Mokka Fiord in summer 1973, and a projected growth curve from 20 June to 8 August 1973. 12 THE CANADIAN FIELD-NATURALIST Vol. 91 TABLE 2—Mean weights, in grams, of organs from arctic hares (Lepus arcticus) collected on Axel Heiberg Island, Northwest Territories in summer 1973 and winter 1975 (standard deviation in parentheses) Number of rF Collection Sex and eee Heart Lungs Liver piles Stomach period age (n) Right Left (full) Summer 1973 Adult males 10 50.5 60.6 93.7 13-8) 13.5 156.6 (6.0) (20.7) (14.0) (1.1) (1.0) (19.2) Adult females 15 50.0 62.4 118.6 16.7 16.5 188.1 (5.3) (12.5) (23.8) (2.0) (2.2) (67.4) Winter 1975 Adult males 7 44.8 67.2 86.3 12.3 12.0 PNS)22 (4.4) (10.7) (33.6) (1.6) (1.3) (29.8) Adult females 5 41.7 67.5 77.0 11.2 11.1 200.5 (8.3) (24.2) (28.8) (1.8) (1.8) (25.6) *Value of m may vary among organs owing to damage during collection. **Weight of kidney minus fat. number of corpora lutea for left ovaries was 3.3 (SD = 1.5) and for right ovaries 3.2 (SD = 0.4). The difference was not significant. The mini- mum and maximum number of corpora lutea for both ovaries was 5 and !1 respectively. A corpus luteum of pregnancy from a lactating female collected on 19 July is shown in Figure 3. The ovaries of seven females collected from 23 March to 4 May 1975 were examined for developing Graafian follicles and corpora lutea. The ovaries of one of four specimens collected on 23 March contained numerous developing follicles although the ovaries from the other three contained but few. Developing and mature Graafian follicles and corpora lutea atretica were numerous in the ovaries of two specimens collected on 4 April (Figure 4). The right ovary of a female collected 4 May contained five recently formed corpora lutea (Figure 5), the left Ovary two, suggesting that ovulation had oc- curred about | May. The ovaries from the one female collected 4 May 1975 were much larger than ovaries of all other specimens. Corpora lutea in ovaries from the 1973 collection measured approximately 2.5 X 1.5 X 3.0 mm; those from the one speci- men at ovulation in 1975 approximately 2.5X 3.0 5.0mm. Although corpora lutea were easily identified in the ovaries of females collected in the summer of 1973, the structures had disappeared from ovaries collected in March to May 1975. The first young hare was seen on our arrival at Mokka Fiord on 22 June 1973. The brownish- gray coloration of young hares and their habit of remaining motionless behind stones made their detection difficult. Food Habits Thirty-five species or genera of vascular plants were identified in 23 stomach samples of adult arctic hares collected at Mokka Fiord during summer 1973 (Table 3). Of the 35 species or genera of plants recorded, 16 were grasses, 13 were dicots, and 6 were sedges. The mean per- centages of epidermal fragments of those classes in the stomach samples were as follows: dicots 53, grasses 41, and sedges 6. Arctic willow was the most common plant species in summer stomach samples, followed by Dryas integrifolia and the grasses Puccinellia Andersonii, Agropy- ron violaceum, and Arctagrostis latifolia. Those five species accounted for 68.5% of the total plant fragments recorded. Only 11.6% of the recorded fragments could not be identified to genera or species, but all fragments could be classified as being grasses, sedges, or dicots. Twenty-two species or genera of vascular plants were identified in 16 stomach samples of adult arctic hares collected at Mokka Fiord during late winter 1975 (Table 4). Non-woody material made up only 9.2% (SD = 2.9%) by weight of the winter stomach samples. The remaining 90.8% consisted of woody material PARKER: ARCTIC HARE ON AXEL HEIBERG ISLAND 13 FIGURE 3. A corpus lut- eum of pregnancy from a lactating arctic hare collec- ted at Mokka Fiord, Axel Hei- berg Island, 19 July 1973. GLC = granulosa lutein cells; TLC =the- cal lutein cells. FIGURE 4. Developing Graafian follicles in the ovary of an arctic hare col- lected at Mokka Fiord on 4 April 1975. AF = atre- tic follicle; GF = growing follicle. Figure 5. A_ recently formed corpus luteum of preg- nancy of anarctic hare collected at Mokka Fiord on 4 May 1975. 14 THE CANADIAN FIELD-NATURALIST TABLE 3—Plant species composition of stomach contents of 23 arctic hares (Lepus arcticus) shot in the summer at Mokka Fiord, Axel Heiberg Island, Northwest Territories. Deter- minations from microscopic identification of epidermal fragments : Mean Plant species Frequency rage density + SE* Dryas integrifolia 100 16.6+ 2.44 Salix arctica 96 AXVS) se DS Puccinellia Andersonii 96 16.0+ 2.36 Agropyron violaceum 91 NOGs2)) 255) Arctagrostis latifolia 61 45+ 2.09 Colpodium Vahlianum 61 11+ 0.33 Alopecurus alpinus 61 19+ 0.58 Oxyria digyna 57 29+ 1.34 Draba spp. 57 hiss, O29 Taraxacum spp. Sy7/ 13+ 0.38 Carex stans 48 2.0+ 0.56 Eriophorum spp. 39 10+ 0.48 Puccinellia angustata 35 Ail se OF Carex spp. 26 Pgsyae > || CVAl Saxifraga oppositifolia 22 10+ 0.50 Pedicularis spp. 22 0.6+ 0.31 Ranunculus pedatifidus 17 0.3+ 0.19 Poa spp. 17 0.1+< 0.10 Saxifraga tricuspidata 13 0.1+< 0.10 Polygonum viviparum 13 0.1+< 0.10 Papaver radicatum 9 0.1+< 0.10 Ranunculus spp. 9 <0.1+ £0.21 Puccinellia vaginata 9 0.2+ 0.21 Deschampsia brevifolia 9 <0.1+< 0.10 Luzula nivalis 9 01+ 0.13 Festuca baffinensis 9 0.1+< 0.10 Poa arctica 9 <0.1+< 0.10 Epilobium latifolium 9 0.5+ 0.40 Hierochloe alpina 4 <0.1+< 0.10 Poa glauca 4 <0.1+< 0.10 Pleuropogon Sabinei 4 <0.1+< 0.10 Luzula arctica 4 <0.1£< 0.10 Carex nardina 4 (ise) O)A8} Puccinellia poacea 4 <0.1+<0.10 Festuca brachyphylla 4 <0.1 =< 0.10 Unidentified dicots 96 U7) 35) MNe8ks Unidentified grasses 96 3.5+ 0.44 Unidentified sedges 52 0.9+ 0.23 *Relative density = number of recognized fragments of a species expressed as a percentage of the total number of fragments of all species. believed to be almost totally arctic willow. Willow also comprised 15.1% of the non-woody material in the winter stomach contents. Dryas integrifolia, Salix arctica, Puccinellia spp., Agropyron violaceum, and Colpodium Vahlia- num together made up 62.6% of the fragments in the non-woody portion of the winter stomach Vol. 91 contents. The approximate mean percentages of dicots, grasses, and sedges among the non- woody fragments were 70, 21, and 8 respectively. Parasites Cursory examination of the carcasses showed no evidence of internal parasites or pathology. Two adult hares collected in the late winter of 1975 were infested with the flea Hoplopsyllus glacialis glacialis, which has been reported from arctic hares in other regions of northern Canada, but these were the first records for Axel Heiberg Island (G.P. Holland, personal communication 1975). Seventeen specimens of the parasite were found on an adult female hare and two on an adult male. The female had scratched off patches of fur and cut the exposed skin with her claws. TABLE 4— Plant species composition of non-woody stomach contents of 16 arctic hares shot in the winter at Mokka Fiord, Axel Heiberg Island, Northwest Territories. Determinations from microscopic identification of epidermal fragments : Mean Plant species Frequency errs density + SE* Dryas integrifolia 100 37.3 + 3.23 Salix arctica 94 15.1 + 2.10 Puccinellia spp. 81 6.3 + 1.58 Agropyron violaceum 81 2.2 + 0.68 Colpodium Vahlianum 75 1.7+0.35 Draba spp. 63 3.1 + 0.98 Oxyria digyna 63 2.3 + 0.63 Eriophorum spp. 56 1.1 +0.38 Carex spp. 50 1.2+0.53 Arctagrostis latifolia 44 1.3 +£0.58 Alopecurus alpinus 44 0.6 + 0.20 Pedicularis spp. 38 0.4+0.15 Taraxacum spp. 38 0.7 + 0.43 Festuca baffinensis 31 0.6 + 0.28 Carex stans 25 0.6 + 0.33 Polygonum viviparum 19 0.5 + 0.38 Hierochloe alpina 13 0.1 + 0.08 Poa spp. 13 0.2+0.15 Ranunculus spp. 6 <0.1 £0.05 Festuca brachyphylla 6 <0.1 £0.05 Saxifraga tricuspidata 6 < 0.1 + 0.05 Pleuropogon Sabinei 6 <0.1 £0.05 Unidentified dicots 100 10.1 = 0.98 Unidentified grasses 100 8.0 + 0.98 Unidentified sedges 81 4.5+0.80 Unidentified monocots 19 0.8 + 0.43 *Relative density = number of recognized fragments of a species expressed as a percentage of the total number of fragments of all species. 1977 Behavior Adult hares had constructed numerous “scoops” or shallow dish-like depressions in the gravel ridges and slopes, usually facing south or southwest and behind, or sheltered by, a large rock. Hares often used those protected scoops during periods of unfavorable weather. Whether young were born in those scoops 1s not known but that seems likely. The young rapidly disperse, however, to spend the first two weeks of life hiding behind rocks, showing themselves only to nibble on vegetation, or to nurse. Lactating females were scattered throughout the lowlands, the young normally restricting their distribution to the gravel ridges and slopes. By the third week of life, young hares had assumed their gray summer pelage and were forming nursery bands numbering up to 20 animals. Although adult females were still in the area, and occasionally within a herd of young, nursing was not observed. Young hares collected on 12 July were feeding mainly on vegetation although hardened chunks of milk remained in the stomach. The herds of young hares con- tinued to grow in size throughout July. By early August most hares at Mokka Fiord had formed into several large herds, containing both young and adults, and were moving erratically throughout the valley. The behavior of hares to human approach varied. Sudden approach, however, usually brought the hare to its hind legs, standing erect on its toes. That position affords the hare a better view and is probably the reason for its use. If the hare is unable to distinguish the object of disturbance and a rapid approach is continued, it usually flees in “kangaroo fashion.” If pur- sued, the hare quickly drops to a quadrupedal position and moves up slope to cover, often assuming an upright stance once high ground is reached, to look back at the object of danger. Solitary lactating females showed little fear of humans, often allowing an approach of several metres or less before slowly hopping away. Herds of young hares in late July and August exhibited a more nervous behavior, often fleeing for a kilometre or more on first sight of a human. I believe heavy predation on young hares by wolves (Canis lupus) and arctic fox (Alopex lagopus) caused that extreme flight behavior. Once the hares identified their pursuer as being PARKER: ARCTIC HARE ON AXEL HEIBERG ISLAND 15 neither wolf nor fox, however, one was able to walk amidst them and be completely ignored. The behavior of the herd in winter was similar to that of late summer. Solitary hares in winter generally displayed more fear than in summer, but again individual behavior varied. The two major predators of the hare at Mokka Fiord were the wolf and arctic fox. Killing was not observed, but both predators are believed to have relied heavily on young hares in the summer of 1973. Arctic fox were often seen carrying young hares to a nearby den. When young hares approached 2 kg in August, and became difficult for a fox to carry intact, the latter often transported the hare carcass in halves. The remains of a young hare, approximately 10 days of age, was found at the nest of a Snowy Owl (Nyctea scandiaca) in 1973. The scarcity of owls in the study area, however, diminishes it as an important predator of the arctic hare. Discussion Arctic hares at Mokka Fiord, Axel Heiberg Island, are heavier than most southern races of the species but comparable in weight to speci- mens from other high arctic locations (Howell 1936) and Newfoundland (3.4 to 4.9 kg) (North- cott 1974). Waterston and Waterston (Water- son, G. and I. Waterson. 1972. Report on wildlife, vegetation and environmental values on the Queen Elizabeth Islands (Ellesmere and part of Axel Heiberg) June 28-August 15. Type- written report for Canadian Wildlife Service, Ottawa) recorded the weight of an adult lactat- ing female on 15 July 1972 at Fosheim Penin- sula, Ellesmere Island as 9 1b (4.1 kg). The mountain hare of Scotland (Flux 1970) and Norway (Walhovd 1965), however, weighs less than Lepus arcticus monstrabilis. The tendency for females to be generally larger in body size and weight than males is also in agreement with studies on hares in Scotland (Flux 1970) and Norway (Walhovd 1965). The decrease in weight by both sexes between summer and winter samples reflects a general summer-to-winter weight loss apparently typical of all species of hares (Hewson 1968). The weight loss is believed to result from a decrease in the quality of diet and approach of the breeding season, although*‘the influence of juveniles in the 16 THE CANADIAN FIELD-NATURALIST winter sample cannot be ignored. The rate of growth of young arctic hares is much greater than for young mountain hares. Both Hewson (1968) and Flux (1970) report a weight gain of 14 g/day for mountain hares under 2 kg. Arctic hares during their first month of life exhibited an average daily weight gain of 45 to 50 g/day. Flux (1971) refers to a 10% seasonal loss in kidney weight of European brown hares (Lepus europaeus) in New Zealand. I found kidney weights of arctic hares in winter to be 11-12% lower for males and 33% lower for females than summer weights. Dauphiné (1975) noted a greater loss in the weight of kidneys from female than from male barrenground caribou (Rangifer tarandus groenlandicus) in northern Canada, although the difference was not as great as that reported here for the arctic hare. The loss of kidney weight during the winter period, as well as from the other major organs of arctic hares, is probably, as suggested for red deer (Cervus elaphus) by Batcheler and Clarke (1970), a physiological response to a decrease in forage quality and quantity and a reduction in the basal metabolism. Arctic hares on Axel Heiberg Island have one litter per year, as do all populations of hares north of the treeline. In Norway the mountain hare may have up to three litters per year, averaging two to four young each litter (Wal- hovd 1965), as does the mountain hare of Scotland (Flux 1970). In Newfoundland the arctic hare may produce two to three litters per year (Northcott 1974). The Alaskan hare is similar to the arctic hare in having only one litter per year (Walkinshaw 1947). A decrease in the number of litters per year appears to be offset by an increase in the number of young per litter. A corpora lutea count of 6.5 suggests the average litter size for hares on Axel Heiberg Island is approximately 5(R. D. Baker, personal communication), slightly less than earlier re- cords of 6 to 7 for hares from Greenland (Pedersen 1926; Manniche 1910, cited by Howell 1936), and 7 to 8 reported for northern Ellesmere Island (Feilden 1877, cited by Howell 1936). On Brunette Island, Newfoundland, J. Inder (per- sonal communication 1976) reported a mean litter size for nine arctic hares of 3.8, the range being from 3 to S. Vol. 91 I first saw young arctic hares at Mokka Fiord on 22 June 1973. Waterston and Waterston (ibid.) reported that female hares on the Fosheim Peninsula, Ellesmere Island gave birth to young between 29 June and | July 1972. On Greenland, young hares were first reported on 10 June (Walkinshaw 1947). In the Canadian High Arctic it appears that most births of arctic hares occur in the last 10 days of June. Examination of ovaries from the 1975 collec- tion suggests ovulation occurred approximately 1 May, giving a gestation period of 53 days. If we assume the birth of hares probably occurred several days prior to our arrival, the gestation period would be approximately 50 days, similar to the period of gestation of 50.3 + 1.3 days for mountain hares in northern Scotland (Flux 1970). Ovarian sections from females collected in the late winter and spring of 1975 suggest that an increase in follicular development begins near 1 April and continues throughout the month. Copulation induces ovulation in hares and the recently formed corpora lutea in the ovaries of a hare collected on 4 May suggests breeding took place near | May. Although structures resem- bling old corpora lutea of pregnancy were present in the ovaries of several females collected in late winter 1975, determining breeding success of the previous year by acount of such structures was not attempted and is not recommended. Arctic hares showed a wide selection in their summer feeding habits, although willow, Dryas integrifolia, and several species of grasses made up the bulk of their diet. Whereas the new green growth of grasses and forbs composed the bulk of their summer diet, woody material accounted for over 90% of the winter diet, most of which is believed to be arctic willow. Approximately 15% of the herbaceous component of the winter diet was also willow, suggesting the total proportion of willow in the winter diet may approach 95%. Sedges were not important in the diet of hares at Mokka Fiord. Sedges are largely confined to the low-lying meadows, areas where hares were seldom seen in summer or winter. The elevated and dry gravel slopes, where hares were most often found at all seasons, supported a sparse but rather diverse flora. Willow, Dryas integri- folia, and a variety of grasses and forbs were the dominant vegetation, suggesting that availa- 1977 bility rather than selectivity probably dictates the diet of the arctic hare. I believe hares utilized the slopes more for escape and food availability because of a light snow cover, rather than food selectivity. Information on the feeding habits of arctic hares in northern Canada is all but absent in the literature. Studies of the mountain hare in Sweden (Lindlof et al. 1974), northeast Lapland (Pullianinen 1972), and Finland (Nyholm 1968) stress the importance of willow in the winter diet. Winter habitat of hare in Scandinavia, however, often includes regions which support shrubs and trees. Bonnyman (1975), after observing feeding hares in late May, mentions the importance of willow in the winter diet of hares on Fosheim Peninsula, Ellesmere Island. Arctic willow is the dominant plant species over much of the Fosheim Peninsula and on eastern Axel Heiberg Island (Inglis, J. T. and C.J. Jonkel. 1972. Ellesmere Island range studies. Canadian Wildlife Service, Ottawa, MS report. 24 pp.). The abundance of that food source, a light covering of winter snow assuring continuous availability, and broken terrain providing escape cover, may explain the abun- dance of hares in those areas in contrast to their relative scarcity over most of the Canadian Arctic. My observations suggest that the arctic hare may be an important competitor with muskoxen and caribou during the winter period when all three species feed upon willow. Whether such competition reaches a level where one or more of the mammal species is adversely affected has not been demonstrated. Such competition would occur only where arctic hares reach extremely high densities such as on Axel Heiberg and Elles- mere Islands. My observations indicate that caribou and muskoxen are not as species- restricted in their winter diet as the hares. Caribou feed extensively on upland grasses and sedges while the winter diet of muskoxen may contain a high proportion of lowland sedges. At Mokka Fiord, one herd of 250-300 hares spent the winter of 1974-75 within an area of approximately 35 km2. Their winter movements could be traced in March by distribution of pellets, remains of broken and uprooted willow, and trampled snow. An area used by hares in early winter becomes virtually unavailable for PARKER: ARCTIC HARE ON AXEL HEIBERG ISLAND 17 further use by hares, muskoxen, or caribou. The loose snow cover is quickly compacted by the hares, and the extreme temperature and wind abrasion create a firm layer which adheres to the ground. Although perhaps only a few centi- metres thick, it becomes extremely hard. In contrast, adjacent areas not trampled by hares build up a deeper covering of snow; a typical late winter profile would show a thick upper crust over a layer of loose ice granules and air pockets. Where the snow has not been disturbed, a muskox can expose an area of vegetation with several blows of its front feet, tossing the broken crust aside and removing the loose ice granules with its hoof or muzzle. Althougha potential for interspecific competition for food appears to exist, muskoxen at Mokka Fiord displayed such a broad selectivity of habitat for feeding that adverse effects of competition were negligible or absent. A high density of hares must certainly play an important role in the energy-flow cycle in a relatively closed high-arctic ecosystem such as the Mokka Fiord valley. The importance of muskoxen, caribou, and hares in sucha system 1s poorly understood. A systems approach for further research is highly recommended for such unique areas as the Mokka Fiord valley and Fosheim Peninsula on Ellesmere Island. Acknowledgments I am grateful to F. Brazeau and R. K. Ross, Canadian Wildlife Service, for their assistance during the collection of arctic hares; to B. Campbell, National Museums of Canada, Ottawa, and J. Maxwell and D. Morton, Cana- dian Wildlife Service, Sackville, for laboratory preparation and analysis of specimen material; and to G. P. Holland, Department of Agricul- ture, Ottawa, for the identification of ecto- parasites. I also extend my appreciation to J. Inder, Newfoundland Wildlife Service, St. John’s, and D. Flook, Canadian Wildlife Service, Ottawa, for helpful suggestions and criticisms of the completed manuscript. Literature Cited Anderson, R. M. 1965. Methods of collecting and preserv- ing vertebrate animals. National Museum of Canada Bulletin Number 69. 199 pp. Banfield, A. W.F. 1961. A revision of the reindeer and 18 THE CANADIAN FIELD-NATURALIST caribou, genus Rangifer. National Museum of Canada Bulletin Number 177. 137 pp. Batcheler, C.L. and C.M.H. Clarke. 1970. Note on kidney weights and kidney fat index. New Zealand Journal of Science 13(4): 663-668. Bergerud, A. T. 1967. The distribution and abundance of Arctic hares in Newfoundland. Canadian Field-Naturalist 81(4): 242-248. Bonnyman, S.G. 1975. Behavioural ecology of Lepus arcticus. M.Sc. thesis, Carleton University, Ottawa. 35 pp. Dauphiné, T.C., Jr. 1975. Kidney weight fluctuations affecting the kidney fat index in caribou. Journal of Wild- life Management 39(2): 379-386. Feilden, H. W. 1877. On the mammalia of North Green- land and Grinnell Land. Zoology 1(8): 313-321; (9): 353-361. (Not seen by author.) Flux, J. E.C. 1970. Life history of the Mountain hare (Lepus timidus scoticus) in north-east Scotland. Journal of Zoology (London) 161: 75-123. Flux, J. E.C. 1971. Validity of the kidney fat index for estimating the condition of hares: a discussion. New Zealand Journal of Science 14(2): 238-244. Handley, C. O., Jr. 1952. A new hare (Lepus arcticus) from northern Canada. Proceedings of the Biological Society of Washington 65: 199-200. Hewson, R. 1968. Weights and growth rates in the moun- tain hare Lepus timidus scoticus. Journal of Zoology (London) 154: 249-262. Howell, A. H. 1936. A revision of the American Arctic hares. Journal of Mammalogy 17(4): 315-337. Lindlof, B., E. Lindstrom, and A. Pehrson. 1974. On activity, habitat selection and diet of the mountain hare (Lepus timidus L.) in winter. Viltrevy (Stockholm) 9(2): 27-43. Manniche, A. L. V. 1910. The terrestrial mammals and Vol. 91 birds of north-east Greenland. Meddelelser om Grdnland 45: 1-200. (Not seen by author.) Nelson, E. W. 1934. New subspecies of the American Arctic hare. Proceedings of Biological Society of Wash- ington 47: 83-86. Northcott, T. H. 1974. The land mammals of insular New- foundland. Wildlife Division, Newfoundland Department of Tourism. 90 pp. Nyholm, E.S. 1968. Ecological observations on the rabbit on Krunni Islands in Ii and in Kuusamo. Suomen Riista 20: 15-31. Pedersen, A. 1926. Beitrage zur Kentniss der Saugetier und Vogelfauna der ostkuste Grdlands. Meddelelser om Grgnland 68: 151-249. (Not seen by author.) Pullianinen, E. 1972. Nutrition of the Arctic hare (Lepus timidus) in northeastern Lapland. Annales Zoologici Fennici 9: 17-22. Tener, J.S. 1954. Three observations of predators attack- ing prey. Number 2, Polar wolf attacking arctic hare. Canadian Field-Naturalist 68(4): 181-182. Thompson, H.A. 1967. The climate of the Canadian Arctic. In The Canada Year Book, 1967. Dominion Bureau of Statistics, Ottawa. pp. 55-74. Walhovd, H. 1965. Age determination of the hare (Lepus timidus Linne) with data on age and sex ratios, growth and body weight. Papers of the Norwegian State Game Research Institute, Series 2, Number 22. 57 pp. Walkinshaw, L. H. 1947. Notes on the Arctic Hare. Jour- nal of Mammalogy 28(4): 353-357. Watson, A. 1954. Observations on some mammals in Cumberland Peninsula, Baffin Island, in 1953. Canadian Field-Naturalist 68(4): 56-60. Received 23 April 1976 Accepted 11 August 1976 Life History Observations of Three Species of Snakes in Manitoba PATRICK T. GREGORY Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Present Address: Department of Biology, University of Victoria, Victoria, British Columbia V8W 2Y2 Gregory, Patrick T. 1977. Life history observations of three species of snakes in Manitoba. Canadian Field-Naturalist 91(1): 19-27. Abstract. Since little is known about the natural history of plains garter snakes, red-bellied snakes, and green snakes, especially in the northern parts of their ranges, various data on these three species were collected over a 34-year period in the Interlake region of Manitoba. As the ecology of the red-sided garter snake in this area is considerably different from elsewhere in its range, similar geographic variation might be expected for the three species studied here. Few differences are in fact obvious; diets, activity cycles, growth, and reproductive features are all basically similar to those in other parts of the species’ ranges. A comparison of the two garter snake species would be of interest for future study because in Manitoba these species apparently exhibit similar growth and reproductive potential, as they do in other parts of their joint range, but have rather different annual cycles of activity. Considerable information has been published recently concerning the natural history of the red-sided garter snake (Thamnophis sirtalis parietalis) in Manitoba. These publications have covered activity both at overwintering sites (Aleksiuk and Gregory 1974; Gregory 1974) and in summer habitat (Gregory 1975; Gregory and Stewart 1975) and have concentrated on the Interlake region of the province, between Lake Manitoba and Lake Winnipeg. To date, how- ever, little work has been done on the other four species of snakes which occur in Manitoba: the western plains garter snake (Thamnophis radix haydeni), the northern red-bellied snake (Store- ria occipitomaculata occipitomaculata), the smooth green snake (Opheodrys vernalis), and the plains hognose snake (Heterodon nasicus nasicus). The subspecific status of Opheodrys vernalis in Manitoba is unclear; Grobman (1941) considers them to be vernalis X blanchardi. Thamnophis radix, Storeria, and Opheodrys all occur in the southern part of the Interlake region, but none 1s apparently as abundant as T. sirtalis in this area. In all three cases, the southern Interlake is the northern limit of distribution for the species at that longitude (Logier and Toner 1961; Conant 1975). Tham- nophis radix, however, does range somewhat further north to the west of the Interlake region, closely following the limit, of prairie grassland. Jordan (1967) indicated that this species is largely restricted to this habitat. Heterodon is confined to the southwestern portion of the province (Scott 1970). While conducting field studies of T. sirtalis from fall 1969 to spring 1973, I occasionally encountered individuals of T. radix, Storeria, and Opheodrys. Since little is known about the ecology of these species, especially near the northern limits of their ranges where interesting life history strategies might be expected, I decided that it would be worthwhile to collect information on various aspects of their natural histories. Data on seasonal activity, diets, body size and age classes, and various aspects of reproduction were of particular interest as they could be readily obtained from living animals. A few attempts were made to obtain information on hognose snakes, but only one road-killed specimen was found. Specimens of Heterodon were occasionally brought in to the Department of Zoology, University of Manitoba, the most notable being a juvenile found as late as | October in 1972, but no data were obtained from any of these. Study Area and Methods Most of the data presented in this paper were collected from snakes found in the southern Interlake region of Manitoba. This area, des- cribed in an earlier paper (Gregory and Stewart 1975), consists of a series of ridges covered with 19 20 THE CANADIAN FIELD-NATURALIST aspen forest and cleared farmland with marshes in the depressions between the ridges. Snakes were occasionally found in other parts of Manitoba such as around the southern part of Lake Manitoba or in the southwestern portion of the province. Both of these areas are generally more open, with considerably more grassland, than the Interlake. No particular sampling plan was followed with respect to the collection of data as the snakes were generally found while I was search- ing for T. sirtalis. Also, because I did not decide to collect data on the three species in an organized way until I was about half-way through my studies, the data are rather incom- plete, varying from simple reported sightings to full information on particular specimens. Sampling effort was fairly evenly spread over the whole study period. In April, May, September, and October my efforts were concentrated at hibernacula of 7. sirtalis; in June, July, and August I generally searched for snakes away from known denning areas. The time period covered by this paper is from the autumn of 1969 to the autumn of 1972. In general, captured snakes were sexed, measured (snout-vent length, SVL) to the nearest 5 mm, palpated to force regurgitation of stomach contents, and individually marked for future identification by the removal of subcaudal scutes in unique combinations. The reproductive con- dition of females was noted and some gravid females were taken back to the laboratory where further reproductive information (dates of birth, brood sizes, etc.) was collected. Results Thamnophis radix, Plains Garter Snake During the course of this study, 56+ specimens of T. radix were collected in spring and fall at communal hibernacula largely occupied by T. sirtalis, more than 51 specimens were observed and/or collected at locations other than dens. Earliest and latest dates of observation were 26 April (1971) and 16 October (1970) respectively, both at den-sites. Earliest and latest observa- tions away from dens were made on 2 May (1972) and 4 October (1970). Thirty-three of the snakes found at dens were females and 12 were males; the sexes of the Vol. 91 remaining snakes were not noted. All but seven of the specimens from dens were caught in the fall; three of the seven found in spring were crow- kills. Only three of the snakes marked at dens were recaptured and in each case the recapture was made in the same fall season as that in which the snake was first seen; the longest interval between first and second captures was |1 days. Only one attempt was made to palpate food from a snake’s stomach at a den-site and no food was obtained. Thamnophis radix was occasionally found at all dens numbered in Figure 2 of Gregory and Stewart (1975) and at a few others not indicated in that figure. By far the largest number (22) was observed at Den |. In addition, more specimens (22+) of T. radix were observed at dens in the fall of 1970 than at any other time. Two instances of suspect interspecific mating activity were ob- served at dens in 1972 (8 and 10 May); in both cases, a female 7. radix was being chased or mobbed by a few male T. sirtalis in the manner characteristic of mating T. sirtalis (Aleksiuk and Gregory 1974). The outcome of this activity was not observed either time as the groups rapidly dispersed upon my arrival. Most snakes encountered away from dens in summer were found in or near marshes or in meadows adjacent to marshes; snakes were occasionally found in areas of aspen forest. Road-kills formed the majority of the more than 20 dead specimens found, while more than 31 snakes were caught or observed live. Thirty-five of the snakes sexed were females, and 13 of these were gravid. Seven of the sexed snakes were males. None of the summer-marked snakes was ever recaptured. Of 21 specimens palpated, 14 produced no stomach contents. Three snakes each contained one wood frog ( Rana sylvatica), one a chorus frog (Pseudacris triseriata), one seven unidentified tadpoles, one an unidentified leech, and the last contained unidentifiable remains. Six of the gravid females found in summer yielded information concerning brood sizes, time of birth, and other reproductive features. These data are summarized in Table 1. The three gravid females kept in an outdoor compound at the University of Manitoba (see Table 1) also provided the only individual growth data I was 1977 TABLE 1—Summary of information on brood sizes and birth dates for Thamnophis radix Date found S_V length,mm_ Reproductive information 22 June 690 Kept in outdoor compound (1972) until 31 July. Gave birth to 35 live and 4 stillborn young on 3 August 22 June 710 Kept in outdoor compound (1972) until 25 July. Gave birth to 51 live and 3 stillborn young on 8 August 3 July 645 Kept in outdoor compound (1972) until 31 July. Gave birth to 11 live and 5 stillborn young on 24 or 25 August 3 July 580 Found as road-kill and (1972) dissected; contained 28 embryos and 41 ovulation scars (corpora lutea) 20 July 660 Found as road-kill and (1972) dissected; contained 14 embryos (similar to stage 37 of T. sirtalis (Zehr 1962)) and 33 ovulation scars 11 August 650 Kept in laboratory. (1971) (10 Sept.) Gave birth to 25 live and 1 dead young between 3 and 6 September in- clusive (average SVL of 22 young was 187.3 mm) able to obtain; these data are presented in Table ee The SVLs of all measured snakes are shown in Figure 1; 41 of these were obtained at dens and 22 away from dens. These data are combined for all years, shown by calendar date, and divided GREGORY: LIFE HISTORY OF THREE SNAKE SPECIES, MANITOBA 21 into males, obviously gravid females, other females, and unsexed. The lines enclosing various groups of points indicate what I feel may represent real age classes, based on apparent tendencies of increasing size throughout the summer and taking into account the size of newborn young (Table 1). Storeria occipitomaculata, Red-Bellied Snake No overwintering sites were known for this species in the Interlake. Most specimens were found in generally open areas such as marshes or meadows, although a few were found in areas of aspen forest. Both gray and browncolor morphs were included in the sample, but the number of each was noted. Twenty-eight specimens were found, six of them dead. Eighteen were identi- fied as females, of which eight were obviously gravid, and two were identified as males. None of those marked was ever recaptured. Earliest and latest calendar dates of observation were 31 May (1970, 1972) and 5 October (1971). Seven snakes were palpated without success for stomach contents. No information on brood sizes was obtained, although one female produced at least two young in captivity between 28 July and 3 August in 1972. One road-killed specimen found on 6 August 1971 had six poorly developed embryos in the left oviduct. A second road-kill found on 22 August 1972 had a copulatory plug in the cloaca indicating recent mating. Figure 2 shows the SVLs of the 12 Storeria which were measured in the same manner as 1n Figure | for 7. radix. Opheodrys vernalis, Smooth Green Snake Forty-one green snakes were found during the study period, 11 of them dead. These snakes were found about equally in wooded areas and in TABLE 2—Snout-vent lengths (mm) of three gravid female Thamnophis radix at various dates during period when held at University of Manitoba in 1972. Vertical lines indicate time interval when parturition occurred Snake No. 22 June 3 July 25 July 9 August 29 August l 690 | 705 2 710 730 | 735 3 645 | 660 22 THE CANADIAN FIELD-NATURALIST 800 600 SVL (mm) 200 WORE EOE AS eS) July May June Vol. 91 29 18 7, 27 ah 3 Aug. Sept... Oct: FIGURE |. Distribution of sizes of Thamnophis radix over active season. o = gravid female, e = non-gravid female, x = male, A = unsexed. Points between vertical solid lines represent summer-caught snakes; other points represent den-caught specimens. Enclosed groups of points represent presumed year classes, dashed lines for females and solid lines for males. Arrow connects two points representing same individual. Parenthesized numbers indicate more than one snake of a given size at a particular date. 300 100 SVL (mm) 30 19 9 June July 29 27 Sept. 18 7, Aug. Zé FIGURE 2. Distribution of sizes of Storeria over active season. Symbols as in Figure 1. meadows adjacent to marshes. No denning sites were known for green snakes in the study area. Nineteen of the sexed individuals were females, of which eight were gravid, and 13 were males. No marked snakes were subsequently recap- tured. The earliest date of observation was 28 April (1972) and the latest 23 August (1971). Twenty-three stomach examinations were made. Seven of these produced food items. Three snakes each contained one grasshopper, two contained one caterpillar, one contained two caterpillars, and one contained arthropod 1977 remains. Further taxonomic identification of these prey items was not attempted. Clutch sizes were obtained for four gravid females. One female measuring 285 mm SVL was found on 26 June 1972 and palpated, revealing six ova. Three females found in a group on 26 June 1972 (referred to in Gregory 1975) contained five eggs each. The SVLs of these snakes were 280, 285, and 290 mm. One of the snakes died two days after capture and was found, upon dissection, to contain eggs with shells. A second laid three eggs on 29 June and then died; the remaining two eggs had shells. The third specimen laid her eggs between 14 and 17 July, inclusive. None of the 15 eggs survived to hatching, but the conditions provided them were not optimum. A female caught on 9 July 1972 (275 mm SVL) had very loose skin in the abdominal region, similar to that of females which have recently oviposited. Figure 3 summarizes SVL data for the 24 specimens of Opheodrys which were measured. Discussion Habitats, Food, and Seasonal Activity The habitats described for the three species in this paper are probably not representative since habitat type was not very often noted. In addition, most of my field time was spent in 400 ZOO SVL (mm) 19 June GREGORY: LIFE HISTORY OF THREE SNAKE SPECIES, MANITOBA 23 marshes and adjacent meadows, the usual summer habitat of T. sirtalis (Gregory and Stewart 1975), thus biasing the capture locations of the other three species. Stomach examinations made of all three species suggest a similar situation to that des- cribed for T. sirtalis (Gregory and Stewart 1975): snakes feed fairly infrequently. The stomach samples obtained from T. radix, although few in number, all represent prey items also eaten by T. sirtalis. D. Platt (personal communication) found that the two species had similar diets in Kansas. No samples were obtained from Storeria, but Stebbins (1966) and Conant (1975) indicate that this species eats slugs, earthworms, and soft-bodied insects. These food types are all available in the Interlake. Opheodrys, as indi- cated elsewhere (Blanchard 1932: Judd 1960; Stebbins 1966; Conant 1975), is largely insectiv- orous. Thamnophis_ radix, Storeria occipitoma- culata, and Opheodrys vernalis all appear to have a longer active season away from hiber- nacula each summer than do sympatric popula- tions of T. sirtalis. | have occasionally found T. sirtalis active away from den sites at the times of earliest and latest captures recorded here (the earliest capture for Storeria and latest capture for Opheodrys are ignored here), but T. sirtalis is 7 29 July 18 Aug. 9 FIGURE 3. Distribution of sizes of Opheodrys over active season. Symbols as in Figures | and 2. 24 THE CANADIAN FIELD-NATURALIST generally congregated at denning sites at such times. It is unlikely that the early and late 7. radix, Storeria, and Opheodrys specimens were simply exceptional individuals. The relatively small number of 7: radix caught at dens in spring, as compared to fall, suggests that dispersal in this species must take place shortly after emergence. Also, those specimens found at dens in spring were generally found early in the season. Lang (1971) found that Storeria arrived later in fall and emerged earlier in spring at hibernacula in Minnesota than did either Opheodrys or T. sirtalis (mostly young). In all three species, dispersal took place very soon after spring emergence, although the overall period of emergence was fairly lengthy. By contrast, adult male 7. sirtalis in the Interlake may spend as much as a month at den- sites before dispersing in spring (Gregory 1974), this tenure being apparently related to the intense mating activity characterizing this stage of the annual cycle. Females disperse after mating. Individual snakes of both sexes spend considerable time at the den-sites prior to entering into hibernation in the fall. For some males, the time spent in the vicinity of hiber- nacula may represent half of the annual active season. Storeria and Opheodrys in the Interlake presumably overwinter communally in aban- doned ant mounds, as described by Lang (1969, 1971) in Minnesota and by Criddle (1937) near Treesbank, Manitoba. Being small species, they (as are young-of-year garter snakes) are prob- ably capable of using a large variety of structures in which to hibernate and likely have little difficulty locating suitable hibernacula in their summer ranges. Lang also found young 7. sirtalis in his hibernacula and Criddle found juvenile 7. radix. Young garter snakes of either species are not known normally to overwinter with the adults in the Interlake and possibly also use ant-mound hibernacula. Ant mounds of the type described by the two authors above are abundant throughout my study area but attempts to find some containing snakes failed. Adult 7. radix apparently do not usually hibernate in large groups as do adult 7: sirtalis (Gregory 1973). D. Hart (personal communi- cation), however, has observed one instance of Vol. 91 small-scale communal denning in T. radix. In general, adult garter snakes are probably more restricted in choice of hibernacula because of their relatively large size and either find sites capable of accommodating large numbers (7. sirtalis) or hibernate in smaller numbers (T. radix). Sex Ratios, Growth, and Reproduction The sex ratios presented in this study are biased in favor of females and cannot be considered representative for any of the three species. This is attributable in part to the non- rigorous manner in which data were collected and to the lack of complete information for many specimens. In addition, the relative pro- portions of females for the two live-bearing species (7. radix and Storeria) are further inflated since gravid females are more easily seen and caught. This is because gravid females tend to bask in exposed areas, presumably to pro- mote development of their broods (Blanchard 1937; Gregory 1975). This does not explain, however, why more females than males of 7. radix were caught at dens. Similar samples of 7. sirtalis at the same dens generally yielded more males than females. Thamnophis radix apparently grows at about the same rate as does T. sirtalis, females being larger than males of the same age. This is based mainly on the data presented in Figure | and in Gregory (1973). Similar growth rates are implied for 7. radix in the Chicago region (Seibert and Hagen 1947) where the growing season is from the end of May to the first week in September. This is about the same as the growing season for T. sirtalis in Manitoba. Platt (personal com- munication) found growth rates of T. sirtalis and T. radix to be similar in Kansas, where both species apparently hibernate in their summer habitat. The relatively small amount of growth shown by the captive gravid females (Table 2) may be an artifact of captivity; on the other hand, gravid females of many species of snakes do not feed very often (Gregory and Stewart 1975) so that little growth might be expected even under natural conditions. Communal mating such as described for T. sirtalis (Aleksiuk and Gregory 1974) is not known for any of the three species studied here. 1977 There is in fact no evidence of mating activity of any kind at den sites in these three species. Possibly, however, dispersal in 7. radix, Storeria, and Opheodrys does not involve such long movements as the 4.3 to 17.7 km in T. sirtalis (Gregory and Stewart 1975) and the probability of finding a mate in the summer habitat may be greater. Lang (1969, 1971) in- dicates that Storeria move less than 3000 ft (about 914.4 m) away from their denning sites in summer. The possibility of interbreeding between the two garter snake species is suggested by the observations of male 7. sirtalis chasing female T. radix, although I have never seen what I took to be a hybrid nor have I heard of one. Both these instances were observed near the beginning of the peak period of mating of 7. sirtalis, when the activity is extremely frenzied at den sites. Perhaps male 7. sirtalis are not very discrim- inatory at such times or the female 7. radix had been in close contact with female 7. sirta/is and retained some of their odor. Odor is very important in attracting male TJ. sirtalis to females (Fitch 1965; Aleksiuk and Gregory 1974). It is unfortunate that the end result was not observed in either case, but the lack of known hybrids suggests that nothing viable would have resulted. Both species of garter snake normally mature in their second year (Seibert and Hagen 1947; Fitch 1965; Gregory 1973; Platt, personal communication), although Platt found that in- dividuals of either species could sometimes reproduce in their first year when food was abundant. In the Chicago area, T. radix mates in late April and early May (Cieslak 1945). Birthin T. radix in Manitoba (Figure |, Table 1) takes place from early August on, at about the same time as in 7. sirtalis (Gregory 1973). The one staged group of embryos (Table 1) is in line with development in 7. sirtalis (Gregory 1973), assuming similar stages in both species. Cieslak (1945) found that parturition of 7. radix occurred from 2-23 August, but was concen- trated in the first week of August. In Kansas, 7. sirtalis and T. radix both give birth from late July on (Platt, personal communication). The litter sizes reported for T. radix in Table | appear to be somewhat larger than for female T. sirtalis GREGORY: LIFE HISTORY OF THREE SNAKE SPECIES, MANITOBA 25 of the same SVLs (Gregory 1973), but the sample is small. Platt indicates little difference in litter size between the two species, and Cieslak (1945) reports an average litter size in T. radix similar to that in 7. sirtalis (Fitch 1965; Gregory 1973). The average SVL of the single litter of 7. radix measured (Table 1) is towards the upper end of the range in newborn sympatric 7. sirtalis (Gregory 1973). Accurate estimates of brood size in Storeria were not obtained, but Blanchard (1937) indi- cates a range from | to 13. My data (Figure 2) suggest that young may be born any time from early July on, but so few gravid Storeria were found that this may be very misleading. Blan- chard (1937) indicates that young are generally born from early August on, as in garter snakes. The sizes of gravid females reported here are at the low end of the range or slightly below the range reported by Blanchard, but all of my specimens were collected in early summer before much growth had taken place. Presumably, females are able to bear young by their second year (Blanchard 1937). The discovery of a copulatory plug in a female Storeria in late summer indicates that fall mating takes place, although it may simply be occasional as in 7. sirtalis (Aleksiuk and Gregory 1974). The work of Trapido (1940), however, suggests that fall mating is probably the rule and that the females retain the sperm in utero overwinter. The Storeria found in this study were comparable in size to those measured by Lang (1971). Green snakes in Manitoba are also similar in size to those reported elsewhere (Seibert and Hagen 1947; Judd 1960; Lang 1971), females being larger than males (Seibert and Hagen 1947). They probably begin breeding in their second year (Seibert and Hagen 1947); repro- ductive females in Manitoba are about the same size as those recorded by Judd (1960). Being Oviparous, this species oviposits somewhat earlier in the summer than the other three species bear their young. The evidence is fairly strong (Figure 2, plus other observations) that eggs are laid from late June to mid-July. This is similar to the egg deposition period reported for Opheo- drys near Chicago (Stille 1954). Blanchard (1932), however, found that this species in northern Michigan laid its eggs much later than 26 THE CANADIAN FIELD-NATURALIST this, usually in the first three weeks of August. Judd (1960) reported finding a female which had evidently oviposited very recently on7 August in southern Ontario. This disparity is interesting because Neill (1964) indicates that Opheodrys has a tendency to retain its eggs longer towards the northern part of its range, approaching a state of ovoviviparity. Stille (1954) also notes this phenomenon and Blanchard (1932) alludes to it in citing an example of a clutch which hatched four days after being laid. On this basis, one would expect a later, rather than earlier, Oviposition date for the species in Manitoba. Stille (1954), however, indicates that year-to- year variability in egg deposition dates is fairly great and that early oviposition is correlated with higher average temperatures in May, presumably because development is initiated earlier. Most of my reproductive data were collected in 1972 when May temperatures were much higher than normal (Annual Metero- logical Summary, Environment Canada, Winni- peg 1972); this possibly explains the early dates observed, but more data over several years are obviously needed. Blanchard (1932) indicates that rough handling of the females may cause premature deposition of eggs. The three females from which clutches were obtained were also palpated for food and this may have constituted sufficiently rough handling that premature ovi- position occurred. It is therefore possible that normal oviposition dates may be somewhat later. According to Blanchard (1932), the number of eggs per clutch in smooth green snakes ranges from 3 to 11 with a mode of 7; Stille (1954) and Judd (1960) obtained similar figures. Those clutch sizes reported here are consistent with theirs. Blanchard (1932) also mentions that it is fairly common for female Opheodrys to lay their clutches in two or more batches, although this may often be associated with abnormal con- ditions. This may have been the case with the female that laid three eggs and died before laying the last two. Conclusion Evidently, on the basis of the limited data presented here, the ecology of 7. radix, Storeria, and Opheodrys in the Interlake region of Vol. 91 Manitoba is not greatly different from else- where in their ranges. This is interesting since Interlake populations of 7. sirtalis exhibit many life history features that are considerably dif- ferent from those of populations in other parts of the range (Aleksiuk and Gregory 1974; Gregory 1973, 1974; Gregory and Stewart 1975). Tham- nophis sirtalis is apparently much more abun- dant in my study area than the three species reported on here and perhaps these life history features represent major adaptations which at least partially explain its numerical superiority (Gregory 1973, 1974). The Interlake environ- ment, however, is extremely rigorous for a reptile (Gregory 1973) and further study of the other three species may reveal interesting eco- logical and/or physiological adaptations. A comparison of the life histories of 7. radix and T. sirtalis would be of particular interest since parameters such as annual growth and repro- ductive potential appear to be similar in the two species in Manitoba despite obvious differences in their annual activity cycles. In other areas where the two species occur, they are apparently similar in both respects. Acknowledgments I thank Don Hart (University of Manitoba) for his invaluable field assistance. Both he and Dr. Dwight Platt (Bethel College, North New- ton, Kansas) provided me with valuable per- sonal communications. Two reviewers also made useful comments on the original manu- script. Jeff Lang (University of North Dakota) kindly allowed me access to his M.Sc. thesis. Literature Cited Aleksiuk, M. and P.T. Gregory. 1974. Regulation of seasonal mating behavior in Thamnophis sirtalis parie- talis. Copeia 1974: 681-689. Blanchard, F. N. 1932. Eggs and young of the smooth green snake, Liopeltis vernalis (Harlan). Papers of Michigan Academy of Science, Arts and Letters 17: 493-508. Blanchard, F. N. 1937. Data on the natural history of the red-bellied snake, Storeria occipitomaculata, in Michigan. Copeia 1937: 151-162. Cieslak, E.S. 1945. Relations between the reproductive cycle and the pituitary gland in the snake Thamnophis radix. Physiological Zoology 18: 299-329. Conant, R. 1975. A field guide to reptiles and amphibians of eastern and central North America. 2nd ed. Houghton Mifflin. xvin + 429 pp. 1977 Criddle, R.S. 1937. Snakes from an ant hill. Copeia 1937: 142. Fitch, H.S. 1965. An ecological study of the garter snake Thamnophis sirtalis. University of Kansas Publications, Museum of Natural History 15: 493-564. Gregory, P. T. 1973. Life history parameters of a popula- tion of red-sided garter snakes (Thamnophis sirtalis parietalis) adapted to a rigorous and fluctuating environ- ment. Ph.D. thesis, University of Manitoba, Winnipeg, Manitoba. viii + 98 pp. Gregory, P. T. 1974. Patterns of spring emergence of the red-sided garter snake (Thamnophis sirtalis parietalis) in the Interlake region of Manitoba. Canadian Journal of Zoology 52: 1063-1069. Gregory, P. T. 1975. Aggregations of gravid snakes in Manitoba, Canada. Copeia 1975: 185-186. Gregory, P.T and K.W. Stewart. 1975. Long-distance dispersal and feeding strategy of the red-sided garter snake (Thamnophis sirtalis parietalis) in the Interlake of Manitoba. Canadian Journal of Zoology 53: 238-245. Grobman, A.B. 1941. A contribution to the knowledge of variation in Opheodrys vernalis (Harlan) with the descrip- tion of a new subspecies. Museum of Zoology, University of Michigan, Miscellaneous Publications 50: 1-38. Jordan, O. R. 1967. The occurrence of Thamnophis sirtalis and Thamnophis radix in the prairie-forest ecotone west of Itasca State Park, Minnesota. Herpetologica 23: 303-308. Judd, W. W. 1960. Observations on the habitat, food, reproductive state and intestinal parasites of the smooth green snake at London, Ontario. Canadian Field- Naturalist 74: 100-106. GREGORY: LIFE HISTORY OF THREE SNAKE SPECIES, MANITOBA 27 Lang, J. W. 1969. Hibernation and seasonal movements of Storeria occipitomaculata in northern Minnesota. (Ab- stract.) Journal of Herpetology 3: 196-197. Lang, J. W. 1971. Overwintering of three species of snakes in northwestern Minnesota. M.Sc. thesis, University of North Dakota, Grand Forks, North Dakota. xi + 97 pp. Logier, E. B.S. and G. C. Toner. 1961. Check list of the amphibians and reptiles of Canada and Alaska. Royal Ontario Museum, Life Sciences Division, Contribution Number 53. 92 pp. Neill, W. T. 1964. Viviparity in snakes: some ecological and zoogeographical considerations. American Naturalist 98: 35-55. Scott, V. H. 1970. The western hognose snake in Mani- toba. Zoolog 11(1): 15-19. Seibert, H.C. and C. W. Hagen, Jr. 1947. Studies on a population of snakes in Illinois. Copeia 1947: 6-22. Stebbins, R.C. 1966. A field guide to western reptiles and amphibians. Houghton Mifflin. xiv + 224 pp. Stille, W. T. 1954. Observations on the reproduction and distribution of the green snake, Opheodrys vernalis (Harlan). Natural History Miscellanea, Chicago Academy of Science 127: I-11. Trapido, H. 1940. Mating time and sperm viability in Storeria. Copeia 1940: 107-109. Zehr, D. R. 1962. Stages in the normal development of the common garter snake, Thamnophis sirtalis sirtalis. Copeia 1962(2): 322-329. Received 7 May 1976 Accepted 28 September 1976 Status and Habits of the Cougar in Manitoba ROBERT W. NERO! and ROBERT E. WRIGLEY? \Manitoba Department of Renewable Resources and Transportation Services, 1495 St. James Street, Winnipeg, Manitoba R3H 0W9 2Manitoba Museum of Man and Nature, 190 Rupert Avenue, Winnipeg, Manitoba R3B ON2 Nero, Robert W. and Robert E. Wrigley. 1977. 91(1): 28-40. Abstract. A cougar (Felis concolor missoulensis), collected at Stead, Manitoba in 1973, and 281 well documented sightings establish the species as resident in the province for the first time. Though there are cougar sightings from 1879 to 1975, the majority are recent (40 in 1974). Prior to 1940, the cougar was restricted to the grassland and aspen-oak transition of extreme southwestern Manitoba—within the ranges of mule deer and American elk. After 1940, these two prey species became rare and localized, and white-tailed deer became the dominant big-game animal, spreading far north into the boreal forest. Recent cougar distribution is closely associated with that of the white-tailed deer, an apparent doubling of the cougar’s range. A rough estimate of the cougar population in Manitoba is 50, and it seems likely that some individuals have been observed in Status and habits of the cougar in Manitoba. Canadian Field-Naturalist adjacent Saskatchewan, North Dakota, Minnesota, and Ontario. Until recently, it was generally assumed that the cougar had been exterminated east of the Rocky Mountains, except for a small Florida population of between SO and 100 animals (Wright 1972). Occasional sightings continued, however, in every Canadian province (e.g., Saskatchewan, White 1967; New Brunswick, Wright 1959, 1972) except Prince Edward Island and Newfoundland, and in many central and eastern states (e.g., Pennsylvania, Doutt 1969; Maine to Alabama, Wright 1972). Most reports (from non-biologists) were not taken seriously, and consequently, valuable information that should have been accumulating on this en- dangered species has been lost. The objectives of this paper are to describe the history and present status of the cougar in Manitoba (and to a lesser extent in adjacent regions), relate its distribution in the province to prey and cover, and examine the animal’s habits in this part of its range. Early historical accounts offer little informa- tion on the presence of cougars in Manitoba. The fur trader, Alexander Henry, made no mention of this cat in his extensive travels throughout the southern part of the province and adjacent states during the years 1799 to 1808 (Coues 1897). A possible reference to cougars was the name, Tiger Hills, given by early settlers to a plateau situated south of the Assiniboine River between the Cypress River, Wawanesa, and Ninette. The first accounts of Manitoba mammals by Thompson [Seton] (1886) and Seton (1909) did not include the cougar, but ina subsequent work Seton (1925-1928) listed seven localities in southwestern Manitoba where the animal had been seen or shot. A few additional records found their way into the literature, but despite the availability of a specimen taken just over the border in Saskatchewan in 1948 (Beck 1958), the absence of any authenticated Manitoba speci- men in a museum resulted in this province being omitted from northern distribution limits plot- ted for the species by Young and Goldman (1946) and by Hall and Kelson (1959). In 1973 a cougar was shot in a farmyard at Stead, 56 km northeast of Winnipeg. The speci- men, which is now in the Manitoba Museum of Man and Nature, gives substantial support to the mounting evidence that Manitoba and ad- jacent regions of midwestern Canada and the United States support a resident cougar popula- tion, almost 1000 km east of the recently accepted range for the species. Methods Most of the early records from Manitoba were published by Seton (1925). Thereafter, reports were investigated and recorded by two former directors of the Manitoba Museum, L.T. S. Norris-Elye (1951) and his successor, Richard W. Sutton (1960). W. Harvey Beck, an assistant curator at the Manitoba Museum of Man and 28 Oa Nature, started a card file on cougar records in 1968 to which were added records gathered by Charles H. Buckner, Federal Forest Biology Laboratory in Winnipeg. In 1972 the senior author initiated a program to obtain and record additional past and new cougar observations in order to document further the status of this species in Manitoba. At that time the cougar was not identified as a game species or as a predator and, if anything, was regarded as a rare transient. Letters and a preliminary report were sent to field staff of the Department of Re- newable Resources and Transportation Ser- vices, requesting their cooperation in the project, and a questionnaire was sent to Provincial Game and Fish associations. No attempt was made to advertise broadly our interest in receiving further reports from the general public. Many reports were investigated further by a letter, telephone call, or personal contact with the original observer, in the hope of obtaining date, location, time, lighting conditions, length of observation, distance to animal, color, descrip- tion, behavior, reasons for suspecting cougar, and number of observers. Most observers indicated a reluctance to publicize their obser- vation for fear of inviting ridicule, and it was learned that a number of sightings had been reported to local authorities at an earlier date, but went unrecorded before our survey owing to a general skepticism regarding the occurrence of the cougar in Manitoba. Finally, cougar data were requested from wildlife agencies, museums, and universities in Saskatchewan, Ontario, North Dakota, and Minnesota to compare trends of sightings and the extent of adjacent cougar populations. This study is based on observations by people with a wide range of occupations and back- grounds. How credible are these sightings? Obviously, there is no way to prove the reli- ability of a report short of obtaining a specimen or photograph. Most reports accepted by us as valid and used in this study consisted of a good description of the animal, including its behavior, and pertinent details regarding the circum- stances of the sighting. Our judgment was also based on other factors; for example, when a number of people saw cougars in the same region, unknown to each other, the evidence became more convincing. Other kinds of evi- NERO and WRIGLEY: THE COUGAR IN MANITOBA 29 dence such as tracks, calls, and the presence of partly eaten wild or domestic prey, provided support for some records, but otherwise were not used to establish the occurrence of the species. Knowing that certain other species may be confused with the cougar, namely, deer, wolf, coyote, dog, and lynx, particularly if the obser- vation is made at a great distance or under poor conditions, we made a special effort personally to check reports. Many sightings were rejected on the basis of insufficient evidence. Results Cougar Sightings There are now records of 281 sightings of cougars in Manitoba for the period from 1879 to 1975, four prior to 1900, and including three in which the animal was allegedly killed (Table 1). From 1900 to 1950 there are one to seven reports per decade; however, 10 cougars were supposed- ly killed during those 50 years (Figure 1). Un- fortunately none of these specimens was saved (several were exhibited for a time in local store windows), partly because the provincial museum was not established until 1932. Since 1950 there has been a great increase in the number of reports received, with 159 during the 1970s and 40 for 1974 alone. Numerous sightings have TABLE 1—Number of cougars seen and reported killed in Manitoba from 1879 to 1975 Year Number of Number of sightings kills 1870-1879 | 1 1880-1889 l ] 1890-1899 2 | 1900-1909 7 4 1910-1919 1 | 1920-1929 5 3 1930-1939 6 2 1940-1949 4 0 1950-1959 27 0 1960-1969 68 ] 1970 8 0 1971 14 0 1972 37 0 1973 35 1 1974 40 0 1975 25 0 Totals 281 15 30 THE CANADIAN FIELD-NATURALIST undoubtedly gone unreported from earlier years. Many observers supplied surprisingly good descriptions of the cougar. The following two reports, for example, are typical of many in our files: Murray Thompson, describing an animal he observed in Riding Mountain National Park in midafternoon in August 1972, wrote as follows: “The animal appeared to be tawny brown or light brown, about 4.5 to 5 feet [1.4-1.5 m] long, with a long, long tail like a piece of heavy rope. It had heavy limbs and large paddy feet, and its head appeared about two sizes too small for its body. The animal was seen walking down the road, then it crossed a ditch and disappeared into the bush. It seemed to pay little attention to the car.” In September 1972, RCMP Constable John Ireland noticed a road-killed deer in the ditch while he was on patrol on Highway 59 at Birds Hill Park, just 13 km northeast of Winnipeg. Returning at 0200 hours, he observed a cougar standing over the deer only 5 m away — soclose that the whiskers on the cat’s upper lip were clearly visible. Constable Ireland guessed the weight of the cougar to be about 35 kg. The cat looked into the headlights for several minutes before it ran a few metres, then came back and tried to drag the deer away. After two attempts failed, it ran off into the bush. The following morning the deer was found where it had been dragged 70 m into the bush; it was partly eaten around the rib cage and was covered with leaves — typical cougar signs. By the next day the deer had been all eaten except the vertebral column, head, legs, and skin. Cougars Killed in Manitoba On the night of 25 December 1973, three men investigated a commotion caused by a barking dog and disturbed cattle in the farmyard of William Kowalke at Stead, 56 km northeast of Winnipeg. A large animal was spotted with the flashlight and, believing it to be a wolf intent on the livestock, the men shot what turned out to be an adult cougar. The specimen was eventually obtained by the Manitoba Museum of Man and Nature (Cat. No. 4725), and is the first con- firmed record of the cougar in Manitoba. The animal was a thin, 2-year-old male (aged by M. G. Hornocker, University of Idaho), measur- Vol. 91 ing as follows: total length, 2108 mm; tail length, 800 mm; hind foot, 292 mm; ear, 95 mm; testes length, 25 mm; weight, 43 kg. Male cougars (F. c. missoulensis) from western Canada aver- age 73 kg and 2413 mm in total length (Cowan and Guiguet 1956). There may have been at least 14 other cougars killed in Manitoba since 1879 (Table 1), but none has been substantiated by a museum specimen. A mounted cougar in the J. B. Hales Museum in Brandon may be the animal that was shot 19 km southeast of Brandon in the early 1920s, but this has not yet been proved. The following cougar kills are listed according to date: Pembina Hills (1879); “Plum Coulee in the Souris Country” [probably Plum Creek] (1887); Birtle (about 1895); Makinak (1901-1902); Elphinstone (1904); Brandon (two killed in 1904, skins on exhibit in general store but now lost); Duck Mountain (“early days”); Little Souris River, 19 km SE Brandon (early 1920s); Birtle (1922); Alexander (1926); Pendennis, 15 kmN, 16 km W Brandon (early 1930s, skin exhibited in general store); Souris River Bend area, about 6 km N, 3 km E Margaret (1936-1937); 3 km N, 15 km W Hadashville (1969); 15 km S, 5 km E Stead (1973). Season and Time of Sightings Sightings were recorded from every month, and 220 could be assigned to particular seasons as follows: 40%, summer (June to August); 33%, autumn (September to November); 14%, winter (December to February); 13%, spring (March to May). These results are expected, since summer and autumn are the seasons when people spend the most time travelling on back roads and working on the land. In New Brunswick, Wright (1959) also reported sightings during every month, with 71% observed from June to November, compared to 73% in this study. The exact time of sighting was recorded in 86 cases. Cougars were seen during every hour of the day and night; from 2100 hours to 0600 hours (sun generally at or below the horizon) there were only one to four reports each hour, whereas the peak periods were 1100 hours and 1600 hours (nine reports each) and 0800 hours (seven). There were 26 (32%) reports from 0700 to 1100 hours and 42 (50%) from 1400 to 2000 hours. Cougar activity periods vary according to prey 1977 NERO and WRIGLEY: THE COUGAR IN MANITOBA 31 ie eee pa HS Island Lak} Baie ey Ing e Swan River » Winnipegosis DUCK MIN a fy e Russell RIDING TN e@ McCreary Shoal Lake | % Selkirke ae aN Winnipeg Steinbach S \ ® Morris x Killarne —~e Morden L Emerson FIGURE |. Map of southern Manitoba showing the maximum distribution of mule deer and American elk (solid line) (after Seton 1909), and cougar sightings (dots) and reported kills (stars) prior to 1940. 32 THE CANADIAN FIELD-NATURALIST activity, and although most hunting is done at night, cougars have also been known to kill big game in the evening and midmorning (Seiden- sticker et al. 1973). Sightings of Pairs and Young The cougar may give birth to from one to six cubs at any time of the year, but breeds only once every two or three years. Young remain with the mother until almost full grown at about 2 years of age. Adults are typically solitary and avoid each other, except for short periods of a few weeks when two females may associate or whena male may be courting a female in heat (Seiden- sticker et al. 1973). In Manitoba, sightings of two or more animals at the same time were reported on 14 occasions. Four cases involved pairs of adults, or adults and large young not distinguished as such; three cases of adult(s) and young (two adults and one young, two adults and five young, one adult and one young); three cases of an adult with cubs (twice of one adult and two cubs, one adult with one cub); two cases of two young alone; two cases of cubs alone (four cubs, two cubs); and two instances when single young cougars were observed, one of which was treed by dogs and then shot. Some of these observations lasted many minutes, with cougars running or walking across open fields, jumping over fences, or climbing into trees. Three reports of paired adult(s) and/or young were made independently by different people in the same vicinity and season. Vocalization In the present investigation, loud cougar calls were described in 15 reports, including four while the animal was in full view, and a cougar was seen in the area in eight of the remaining 11 cases. A difference of opinion has existed among cougar experts as to whether the cougar emits a loud scream. Some observers (e.g., Seiden- sticker et al. 1973) admit hearing close-range vocalizations, but never roaring and screaming such as is abundantly described in the literature (Young and Goldman 1946; Barnes 1960). Although calls of bobcats (Lynx rufus) and owls have probably been ascribed incorrectly to cougars On some occasions, there is no doubt Vol. 91 that cougars are capable of producing terrifying calls, since they have been observed screaming in the wild and in captivity. It appears that cougars are rather silent except at mating time (Seton 1925; Wright 1959), and this may partly account for the fact that cougars are heard in one region but not in another. Behavior A peculiar aspect of cougar behavior, pointed out repeatedly in these reports as well as in the literature, is its indifference to being observed from a distance by people on foot or in motor vehicles. Cougars often permitted the approach of a car to within 10 to 100 m before walking leisurely or bounding to the nearest cover. The following case illustrates how bold the cougar may be. On 30 June 1974, at 1400 hours, Tim Sims was cultivating a field near Snowflake, Manitoba, when a cougar came out 400 m from the bush and lay down in a fresh furrow, per- haps to escape flies. Sims watched the animal for an hour as he made several rounds of the field, once passing within 12 m (close enough to see the cougar curl its lips and snarl). It did not appear to be frightened. Sims had seen the same animal in this field 3 days earlier and described it as brown, 0.8 m in height, 2.7 m from nose to tip of tail, tail about 0.6 m with a dark tip. On 8 August Mr. Sim’s son saw a cougar with three cubs on the same land. Food Habits Cougars were seen stalking, chasing, or eating a variety of prey species, and were implicated (by nearby sightings, screams, tracks) in many others. Attacks (prey killed, wounded, or flee- ing) were reported on cattle (13 cases), white- tailed deer (Odocoileus virginianus) (9), horse (3), snowshoe hare (Lepus americanus) (3), chicken and turkey (2), Mallard (Anas platyr- hynchos) (1), hog (2), sheep (1), moose (Alces alces) (1), Formosan deer (1), dog (2), fisher (Martes pennanti) (1), and man (1 possible fatality). Cougars were seen stalking the follow- ing: cattle (3 cases), white-tailed deer (3), beaver (Castor canadensis) (1), pheasants (Phasianus colchicus) (1), and ravens (Corvus corax) sca- venging on a muskrat (Ondatra zibethicus) carcass (1). Cougars were seen scavenging on a white-tailed deer carcass, a trapper’s cache of 1977 carcasses, a trapline (tracks and other sign indicated scavenging ona snared snowshoe hare, and killing and eating a fisher caught in a trap), and in a refuse dump. There were three instances of cougars observed drinking. The Stead cougar had striped skunk ( Mephitis mephitis) hairs in the digestive tract, and porcupine (Erethizon dorsatum) quills in the muscles of the forearms and shoulder regions. Traces of both species are often found in scats of cougars, and porcupines appear to be favored prey (Young and Goldman 1946; Wright 1959; Barnes 1960). One report is of exceptional interest. In the summer of 1972, Walter Larocque herded 32 horses into an old pasture surrounded by bush at The Pas. In an abandoned house on the site, he found moose droppings all over the floor and blood on the door jamb, and concluded the moose had been chased into the house by some animal. One month later he noticed that two horses were missing, and the depredations con- tinued until a total of six mares and one colt had been killed and eaten. The carcasses were found in the bush with all the meat stripped from the bones. It appeared that attacks were launched from a big elm tree that leaned over a path taken by the horses. The horses were herded into a wire corral and when two men went to check them the following morning, they had broken out. The men went to the northend of the pasture and met the horses on the run, followed closely by two tan-colored cougars which came to a halt, their “tongues sticking out of their mouths” from exhaustion of the chase. The larger cat crouched and crawled within 30 m before the men backed away. The horses were gathered and removed from the pasture. Horses, particularly colts, appear to be a favorite prey of the cougar; the large, nearby race of cougar formerly inhabiting North Dakota, F.c. hippolestes, was named for this reputation as a “horse killer.” Because many livestock losses go unreported and government field staff presently lack ex- perience in identifying cougar kills or damage, there may be a larger incidence of cougar preda- tion on livestock than is presently known. There is a single reported case of a cougar stalking people in Manitoba, which ended in tragedy. The following account was published by V. W. Jackson (a University of Manitoba NERO and WRIGLEY: THE COUGAR IN MANITOBA 33 zoologist) in the Winnipeg Tribune (14 February 1942): “Twenty years ago [1922]a mountain lion attacked a boy and a girl at Birtle, Man., killing the latter. When shot it was found to be blind and half-starved.” A search for additional infor- mation on this case, including a local history of the area (Abra 1974), produced no substantiat- ing evidence. Habitat and Distribution of the Cougar and its Prey in Manitoba Remarks on habitat in 255 reports were grouped as follows: 40% “wilderness” (true wilderness and heavily wooded regions with few roads or human habitations), 30% “mixed land” (agricultural lands and towns interspersed with large tracts of forest), and 30% “farmland” (cropland and pasture with forest cover re- stricted to woodlots and river valleys). The pre-1940 records (Figure 1) cover an area of about 87 000 km? and are restricted to the grassland region and the aspen-oak transition (now largely farmland and mixed land) in extreme southwestern Manitoba. From 1941 to 1975 the cougar appears to have extended its range (Figure 2) northward into the boreal forest region and eastward into the Great Lakes - St. Lawrence forest transition (Figure 3). The species now inhabits an area of about 200 000 km? in the southern half of the province, more than twice its apparent former range. Although the cougar accepts an extremely wide variety of prey species (e.g., mice, fish, grasshoppers, small birds, etc.), its residence in an area is still dependent on big game, partic- ularly deer (Barnes 1960). “The range of the panther [cougar] has always coincided with that of the deer, and as the deer have flowed back over the eastern ranges, so has the panther returned to many of its old haunts” (Wright ISS), In early historic times southern Manitoba supported abundant mule deer (Odocoileus hemionus), American elk (Cervus elaphus), moose, caribou ( Rangifer tarandus), pronghorn (Antilocapra americana), and American bison (Bison bison). Rather suprisingly, of these species, only mule deer and elk have been found to play a role in the cougar’s diet. Reported attacks or predation on the other four species are completely absent or restricted to a few isolated 34 THE CANADIAN FIELD-NATURALIST Vol. 91 (>) a)” | }.Nelson House A, j ‘Thompson. a7} j ', ah) ee J oi Za ipinesl Lake Lye y, af Zz G p Cormorant Lake* % t Jor sale Vie i) AD) OO oe TheyPas)& iD Fe Ctr e P i © Swan River | DUCK MIN ! e Russell VG ae N Q McCreary @® @ Neepawa Selkirkef@ e oe e > Portage la Prairie 3 ° QUE, 2 m3 ps) Os 2 7 77 v ° SS 2 p re, = % BS es ay Fe BER ;-—WE — ay ATS Le meee: aa A x wate = —— es fat Ly sug eee —— ZF, SSS vou sek = RATS = — rakes, + = geese se. F =~ LORY z = oan Wa ®, + SA i reeas ( Br ae, - Soff EER wis eeeeeses =. — RES 5 Seaneee : MICH. EEE : EOE 4 _- pee = : ce : (as) Kee SRR Coenen LEER : : : Lee Ogee Sey, : ‘ awe coaee = = : os : : cy : PREETI rene = Ln eon on into pm on tt r : BEE = H 1 eee eee. z ee Og Se La, — 5 — ‘ PALS = : : 4 t : t EEO 4 : : — Bes OH | Q ¥ . os z Dee ae, 7 \ t i omen ge Sas Ss Sm, {9 : Se cos Se l ai —— = Cl CT 4 J MES. = ; rT. oF OWA == —— : = =f a KY. 4 0 50100 200 300 =o oe : ¢ a enn s BP KILOMETERS FIGURE 1. Ontario study location in the north-central portion of the range of C. p. marginata. Map adapted from Conant (1975). Intergradation occurs in zones of overlap. with occasional opportunistic hand sampling. The nets were regularly relocated in a total of 25 sites within the lower sanctuary. Capture by hand was necessary only for terrestrial samples. Carapace total length and carapace curved length were measured with a flexible metal tape rule accurate to 1.0 mm. Weight was recorded by placing the animal on its back in a plastic bucket which was then set on a hanging spring scale precise to 25 g. Sex was determined from the length of the foreclaws. In 1974 each turtle was numbered on the plastron with indelible red ink and when the first animal with an illegible number was recaptured, after approximately 2 months, trapping was discontinued. In the first 3 weeks of June 1975 turtles taken in the nets were marked and released but the main emphasis at that time was observations on shore. In 1975, numbering of individual turtles was accom- plished by notching the marginals with a hacksaw blade (Cagle 1939). Each marginal had been assigned a number so that a notched marginal, or combination of marginals, served to identify turtles in a more permanent manner than the previous year’s ink markings. The three characters used by Hartman (1958) to distinguish C. p. marginata from C. p. picta (Schneider) were measured with calipers exact to 0.01 mm. Alignment of central and lateral laminae was measured and calculated as des- cribed by Hartman (1958) and later by Pough and Pough (1968). One hundred percent dis- alignment describes exact alternation of the laminae as is “typical” in C. p. marginata, while 0% represents the linearly catenated seams “typical” in C. p. picta. The plastral figure, if present, was copied directly onto tracing paper to allow for later experimentation with different plastral figure measurements. Eventually figure length, greatest figure width, longest figure extension along a seam, and plastron length were measured to the nearest 0.5 mm. Figure length and greatest figure width were plotted against plastron length, and longest seam-figure 1977 WHILLANS and CROSSMAN: MIDLAND PAINTED TURTLE, CENTRAL ONTARIO 49 TABLE |—Statistics relating to shell markings of Chrysemys picta marginata in Nogies Creek, Ontario, 1975. n, number in sample; x, mean of the “characteristic” expressed as a percentage for the total of the relevant sample population; SD, standard deviation (one); SE, standard error (one); , population mean estimated by X, subscript 1 represents the male sample, sub- Characteristic Sample composition n x Plastral figure Total sample 47 62.98 length/ plastral Males 222) 64.66 length Females 23 64.13 Greatest plastral Total sample 47 46.85 figure width/ Males 22 43.00 greatest plastral Females 23 50.14 width Longest seam- Total sample 47 20.34 figure-extension/ Males 22 19.10 greatest figure Females 23 22.26 width Light margin/ Total sample 50 1.04 total length Males 23 0.93 Females 26 1.13 'Critical value 95% confidence = 1.960. extension against greatest figure width (see Table 1). Those ratios test the length relation- ships that interested Hartman (1958), the figure width characteristics that concerned Bishop and Schmidt (1931), and the figure extension traits that Carr (1952) found to be important. Hart- man (1958) and Ernst (1970a) indicated a sub- specific differentiation in the width of the anterior light margin on the third lateral. This was measured. The total length (anterior to posterior) of each of the carapace and plastron was recorded. Shell abnormalities were also noted. Observations of nesting activities were con- centrated at four well used sites. A gravel road runs parallel to the west shore of Nogies Creek for a distance of 3.2 km. The turtles had to cross the road to reach three of the nesting locations and were easily seen. Observations started at 0700 hours and ended at dark since there were no references to night activity by C. p. marginatain the literature surveyed. In the beginning the road was traversed every half hour and observations were made on all turtles that could be located from the road. As a result it became obvious that there were daily peak periods of activity and script 2 represents the female sample SD SE 95% confidence Test hypothesis! Mi-uw2 = 0 16.30 2.38 58.19-67.76 8.32 1.77 0.0555 Accept 45.28 9.44 9.16 1.34 44.16-49.54 9.70 2.01 2.7401 Reject 7.58 1.58 8.67 1.26 17.79-22.88 6.66 1.42 1.2951 Accept 9.52 1.98 0.36 0.05 0.94-1.14 0.38 0.08 1.8554 Accept 0.35 0.07 periods of lesser activity. Subsequently activity was monitored every half hour during the known peak periods and at hourly intervals otherwise. The duration of each observation (on one or more than one turtle) varied but the road was not left unmonitored for longer than the intervals given above. The fourth nesting location, having poor accessibility, was visited irregularly. Date, time of day, location, identifying number, and morphological characteristics were recorded for each turtle. When an animal was found to be nesting, its activities were observed from a place of hiding or with binoculars from a distance. Measurements were made after nest- ing, or when the turtle was returning to the water. Nests were measured (diameter of neck, depth of main chamber, depth to bottom of main chamber). The substrate was described, distance from shore estimated, and terrain noted. If a turtle was seen at a nest which was later covered over, the nest was unearthed. From any nest with eggs, clutch size, depth to eggs, and egg size (length and width) were recorded. Air temperature was measured daily with a maximum-minimum thermometer. Daily max- 50 THE CANADIAN FIELD-NATURALIST imum and minimum air temperatures from the nearest official Climatological Station in Cobo- conk were obtained through the Meteorological Applications Branch of Environment Canada. The Corn Heat Units (CHU) were calculated from these (see Brown 1972). Water temperature was not considered because of an incomplete record at the sanctuary; however, the close correlation between air and water temperatures should minimize the importance of this omission (see McCombie 1959). Measurements and Observations Standard Measurements A sample of 50 turtles from Nogies Creek was tested for disalignment of the lateral and central laminae seams. The mean disalignment was 86.4% with a standard deviation of 11.3 and standard error of approximately 1.6. This is in accordance with the findings of Pough and Pough (1968) and Hartman (1958), and is well above the minimum 55% disalignment set by Ernst and Ernst (1971) for the midland painted turtle. Figure 2 shows that of populations studied to date, the Nogies Creek population is “— CAPE COD, MASS. (H 12) Vol. 91 perhaps the closest to expected findings for C. p. marginata populations. The wide range of 70 disalignment percentage points (36-102) is partly explained by one deviant individual (36%). The next value higher than 36% was 73%. Plastral figure measurements are summarized in Table 1. The mean value for figure length as a percentage of the plastral length is 62.9. Greatest plastral figure width averages 46.8% of the greatest plastral width. The mean value for longest seam-figure extension as a percentage of the greatest figure width is 20.3%. Since the Nogies Creek plastral figures were all recorded on tracing paper, they can be compared to the diagrams of C. p. marginata, C. p. bellii (Gray), and an intermediate given by Bishop and Schmidt (1931). A visual inspection of their sketches reveals that in a Nogies Creek sample of 50 turtles the majority (38 or 76%) had what seems to be C. p. marginata plastral figures. None displayed the markings of C. p. bellii, but nine (18%) had intermediate character- istics. No one has yet been able to devise a totally PERCENT 50 CUTTYHUNK, MASS, (P 14) SOUTH AMHERST, MASS. (H 19) NANTUCKET, MASS. (Pp 25) LONG ISLAND, NY. (Pp 73) Northern NEW JERSEY (P 26) DRYDEN, farmpond NY, (H 21) FISH HATCHERY, ITHACA NY. (H 20) FLINT, MICHIGAN (H) SODUS BAY, NY. (H 25) NOGIES CREEK, ONT, (50) P—Pough and Pough (1968) ° | H— Hartman (1958) eo 50 PERCENT FIGURE 2. Percentage of disalignment of central and lateral seams for several populations depicting range, mean, one standard error and one standard deviation on each side of the mean. 1977 reliable statistical method for expressing the differences between the plastral figures of the two subspecies, because of the high degree of individual variation. Bishop and Schmidt (1931) used the width of the plastral figure expressed as a percentage of the plastral width. According to their study, the mean value for C. p. marginata was 36%, for C. p. bellii 74%, and for inter- mediates 55%. The Nogies Creek turtles, with a value of 46.8% (Table 1) lie about halfway between the value for intermediates and that for C. p. marginata. Only eight (16%) individuals had plastral figures with widths less than 36% of the plastral width, and none were in excess of 74%. Three of the Nogies Creek turtles had no plastral figure at all. This characteristic is normal in C. p. picta. The mean value of figure length as a percentage of plastral length for Nogies Creek turtles is 62.9% (Table 1). There are no comparable figures in the literature. Bleakney (1958) indicated two intergrade figure traits: (1) dark spots on the midline, and (2) smaller, often lighter figures, symmetrically distributed about the midline and having blurred edges. None of the Nogies Creek sample dis- played dark spots on the midline and only four turtles were faint in color and had figure lengths WHILLANS and CROSSMAN: MIDLAND PAINTED TURTLE, CENTRAL ONTARIO 51 less than 50% that of the plastron. The longest seam-figure extension was cal- culated as a percentage of the greatest figure width (20.3) with little utility. This relationship did not support the visual impression of the plastral figure as did the aforementioned com- parisons involving figure length and width; however, it has been presented in Table | as the best measure of seam extensions. Other traits compared less successfully to longest seam- figure extension were plastral length, figure length, and greatest plastral width. Hartman (1958) plotted the width of the light margin against carapace length. Nogies Creek painted turtles have been similarly character- ized and compared with Hartman’s results in Figure 3. The Ontario turtles align themselves with the New York samples of C. p. marginata and are clearly distinguishable from the Massa- chusetts intergrade population (C. p. marginata XC. p. picta). All three of the ventral figure relationships, and that between the light margin and carapace length, were tested for sexual dimorphism. The width of the plastral figure, when expressed as a percentage of the width of the plastron, was the only one of these characteristics having two Statistically different sexual values (see Table 1). a S.Amherst, Mass. (Hartman, 1958) © Dryden, N.Y. (Hartman, 1958) @® Nogies Creek, Ont. 5 — E 4 z O a < = 3t a = | <= oe me ca ae oO | x a al a ) = | : 00 0 0) 25 50 75 100 125 150 175 Te eee ea aca| 200 CARAPACE LENGTH (mm) FIGURE 3. Width of the light margin of the third lateral plotted against carapace length. Adapted from Hartman (1958). 52 THE CANADIAN FIELD-NATURALIST These calculations are only slightly significant (test hypothesis value = 2.74, 95% confidence = 1.96) and because of the nonsignificant values for each of the other three shell-marking characteristics, sexual dimorphism was not pursued any further. Other Measurements and Morphometric Obser- vations Frequency polygons for length and weight in the sample of Nogies Creek turtles are given in Figure 4. A total of 195 turtles was captured. They ranged in length from 105 to 181 mm and in weight from 175 to 675 g. According to Ernst (1971a), male turtles in Pennsylvania usually mature at a plastral length of about 70 mm, in their fourth year, and females at approximately 100 mm, in their fifth year. If this is applied to the Ontario population it appears that netting was selective for mature painted turtles, espe- cially since there was no problem sexing even the smallest animals. Females were larger than males. Fifty-one turtles were examined in 1975 for the humped-back condition (“kyphosis”) des- cribed by Ernst (1971b). None displayed the trait. An additional 195 turtles from 1974 were tested for deviation from a straight-line relation- Vol. 91 ship between the total length of the carapace and the curved length of the carapace. No obvious deviants were noted in these Nogies Creek turtles. Ernst (1971b) noted that 0.5% of his sample at the White Oak Bird Sanctuary in Pennsylvania exhibited kyphosis: In a subsample of 51 turtles from Nogies Creek, exterior deformities were recorded: 20 (39%) had deformites of the following kinds. Plastral abnormalities were the most numerous. Ten of the 20 animals (19.5% of 51 sample turtles) had misshaped plastrons. Ernst (1971b) found only one occurrence of this peculiarity in his collection of 929 turtles. In the White Oak study, carapacial abnormalities were found on 6.2% of the deformed turtles. This variation comprised 13.7% of the Nogies Creek sample. Sixteen percent of the 5! Ontario turtles examined displayed unusual appendages. Com- binations of the above deformities were not uncommon in either Ernst’s or the present studies. Spring Movements: Aquatic Figure 5 summarizes the numbers of C. p. marginata captured per net, per day, in 1974. A comparison of the daily air temperatures for the same period of time, also in Figure 5, reveals a CARAPACE TOTAL LENGTH (mm) ! female (s) male (s} 150 160 170 180 = ea NUMBER OF TURTLES ie al Ke 200 250 300 350 400 450 500 550 600 650 700 WEIGHT (g) FIGURE 4. Lengths and weights of male and female C. p. marginata from Nogies Creek. 1977 ----- Max. temperature —— Min. temperature seoeee No. of turtles = ae Data missing (G0) {o) NO (eo) AIR TEMPERATURE (°C) 22 “BS 30 1 5 10 15 WHILLANS and CROSSMAN: MIDLAND PAINTED TURTLE, CENTRAL ONTARIO 53 NUMBER OF TURTLES PER NET 20 . 25 1 5 10 —M AY —e JUNE JULY DATE 1974 FIGURE5. Daily maximum and minimum air temperatures plotted with the numbers of turtles caught per net, calculated per 24-h period in 1974. direct relationship between air temperature and turtle movement reflected in the number cap- tured. But, there often was a time lag between peaks of temperature and activity. There was also a coincidental increase in temperature and movement immediately prior to the initiation of nesting activities. Extensive cloud cover was observed (no data) to have a negative influence on turtle activity. Turtles were first seen nesting on 6 June in 1974 and 12 June in 1975. At the initiation of nesting in 1974 approximately 515 Corn Heat Units (CHU) had accumulated since mid-April (calculated from records at a climatological station 19.3 air km to the northwest). Nesting in 1975 started when 804 CHU had been recorded. In the aquatic netting of 1974, 174 different C. p. marginata were captured, measured, and returned to exactly where caught, before plastral numbering became illegible. Of these, 43 (24.7%) were recaptured, 32 (18.3%) only once, seven (4.0%) twice, two (1.1%) thrice, one 8 times, and one 10 times, for a total of 244 captures. Twenty- six of the 43 recaptured turtles had changed position between captures. In the total of 244 aquatic captures between 22 May and I1 July 1974 there were 159 males and 85 females; the sex ratio was 1.87:1. Because some turtles were captured more than once, actually only 174 individual C. p. marginata were caught in the nets. This included 104 males and 70 females for a sex ratio of 1.49:1. Single captures were recorded for 131 turtles with a 1.34:1 sex ratio. This contrasts sharply with the sex ratio of 2.07:1 for individuals displaying multiple recaptures. When the number of cap- tures per sex are summed, regardless of the number of turtles involved, the sex ratio be- comes 3.67:1. Spring Movements: Terrestrial All 30 terrestrial captures, involving 25 C. p. marginata, were females. Three of the five recap- tures occurred on the day after the initial capture. These three recaptured individuals were all discovered within 5 m of the point of original 54 THE CANADIAN FIELD-NATURALIST capture, one no more than 0.5 m from the first site. Another of the five recaptures was found after 2 days about 15 m from the initial location. The last was relocated 8 days later approxi- mately 150 m from the original site. A large number of empty nest holes was found in the most frequented nesting areas. In 1974 an unrecorded number of unmarked turtles was observed digging nests which they later aban- doned as empty holes. Sometimes these con- tained obstacles such as rock; in others there was no apparent physical restriction. Only three turtles were captured both in the water and on land. One had travelled 250 m from the initial site of netting to an adjacent part of the shore. It was found 100 m from the water by a sandy roadside bank. Two turtles were discovered on land 850 m and 1075m from the original location of capture. These represent the farthest distances travelled by any of the marked animals. The two were found within 30 m of each other at one of the most-used nesting sand banks. One had been previously captured in the water 1075 m to the north along an irregular shoreline. The other was later captured in a net 850 m to the south along a similar shore. From the creek there was no apparent visual clue to the location of the sand bank owing to the high bushes which line most of the water’s edge. Nesting Activities Nesting behavior in painted turtles has pre- viously been described: C. p. marginata (Hart- weg 1944; Carr 1952), C. picta (Babcock 1919), C. p. bellii (Legler 1954; Mahmoud 1968). Al- though direct observations at Nogies Creek largely confirmed these other findings, three additional points were recorded. Four turtles were observed from the time of initial search for a suitable nest site to the final covering over of the eggs. Nesting behavior for an additional dozen animals was noted in part. (1) A female C. p. marginata would move out of the water at a steady pace and with its head level. It would stop upon reaching denser vegetation, lift its head and remain static, or proceed erratically until a new direction was established. This behavior continued until the turtle came to a potential nesting area. Then it would lower its head until almost touching the ground and creep Vol. 91 ahead slowly as if sensing something. Occasion- ally it would scrape the ground with its claws. The site eventually was accepted or rejected. (2) The periodic discharge of liquid from the anus during digging was described by Legler (1954). At Nogies Creek turtles measured prior to laying eggs discharged liquid, but any cap- tured upon completion of a confirmed nesting had no liquid discharge. Turtles continued nesting activities even after discharging their liquid. Thus the liquid, which seemed to aid in digging, is probably not a necessity. (3) When laying was completed, the female turtle would cover the nest hole using its hind legs and tamp the surface down with its plastron. One turtle was observed dragging some nearby (one shell length) loose leaves over the nest with its front legs. Nests in grass were almost in- distinguishable even immediately after their formation. A rain usually obliterated all traces. Seventeen female C. p. marginata were found moving on land and away from the water. They were all discovered between 1500 and 1915 hours (1640 hours mean). The duration of their activities varied from | to 4h with the largest recorded capture being at 2100 hours. Turtles were located anywhere from | to 200 m away from the water on shore (mean 46 m). Approx- imately 37 nests, however, ranged between 2 and 50 m from the shoreline, averaging 20 m. Clutch size in five nests varied from 6 to 9 eggs, with a mean of 7.2. This is within the 3 to 11 egg range listed by Carr (1952). Immediately after laying, the eggs had an average width of 18.5 mm (range 16.9 to 19.5 mm, n= 14) and a mean length of 30.8 mm (range 26.7 to 50.0 mm, n = 14). One nest examined after it was incubated for 73 days contained five apparently healthy eggs (plus four mutilated) which averaged wider but shorter than those newly laid (width: mean 20.5 mm, range 18.1 to 22.2; length: mean 29.0 mm, range 25.5 to 31.1). The comparable ranges listed by Carr (1952) are length 28.6-31.8 mm, and width 17.5—20.6 mm. Mean nest dimensions were as follows: dia- meter of neck, 29.1 mm (n = 3); diameter of main chamber, 65.2mm (n=7); depth to eggs, 25.0mm (n=1); depth to bottom of main chamber, 70.6 mm (n= 6). Ernst and Barbour (1972) described nests of C. picta as having the 1977 following dimensions: main chamber 65-72 mm, neck diameter 41-51 mm, and depth 99-111 mm. Most nests were situated in sandy loam, gravel, or sandy gravel. The ground was clear, or hada grass cover of up to 30 cm in height. All received the sun most of the day and there were few not on level ground or on the lower slopes of an east- facing bank. From the data of this study it is impossible to estimate nesting success. Pred- ators were numerous, racoons, people, and larval insects being common. No vernal emigration as reported by Sexton (1959) was noted in Nogies Creek. Constant water levels regulated by a dam at the lower end of the sanctuary may account for this. Discussion Standard Measurements Bleakney (1958) hypothesized that the north- ward postglacial dispersal of C. p. bellii and C. p. dorsalis (Agassiz) resulted in an intra- specific hybridization around St. Louis, Mis- souri. The subspecies C. p. marginata emerged and subsequently extended its range to its present status. The Nogies Creek turtles clearly display cara- pacial disalignment and light margin width “typical” of C.p. marginata. The plastral figures, however, seem to be wider and perhaps longer than might be expected. This does not necessarily indicate intergrade characteristics of the north-central subspecies with another. It could be evidence of the C. p. bellii influence in Bleakney’s postulated C. p. bellii X C. p. dor- salis origin of C. p. marginata. Similarly, al- though the lack of plastral figure or its reduced size is a trait documented in intergrade C. p. marginata X C. p. picta, its occurrence in this sample might be attributable to C. p. dorsalis ancestry. Plastral figures, however, possibly become fainter with age as all three turtles were large (older?). Masat and Musacchia (1965) found four electrophoretic patterns in blood serum proteins of C. picta. These patterns were randomly distributed among the turtles regard- less of subspecies. Although this does not dispute Bleakney’s theory it does suggest that a more widespread introgression could have in- fluenced C. p. marginata. The plastral figure length and width, in com- parison to the respective length and width of the WHILLANS and CROSSMAN: MIDLAND PAINTED TURTLE, CENTRAL ONTARIO 55 plastron itself, appears to be the most readily quantifiable of the figure characteristics. But, it must be expected that of the subspecific para- meters discussed herein, this will be the most variable as a result of the difficulty in describing the plastral figure. This study clearly demonstrates that even in the north-central part of its range, C. p. marg- inata displays a diversity of individual traits. The necessity of large samples for correctly des- cribing peculiarities in the midland painted turtle is emphasized. It is probable that even in the best of samples, a statistical population of C. p. marginata will only approach the “typical” and defining disalignment, plastral figure dimensions, and light margin widths. The Nogies Creek painted turtle sample was normally distributed with respect to carapace length. Individual turtles ranged in size from 103 to 181 mm. Carr (1952) described the catches of a New York study as consisting of animals ranging from 106 mm to 175 mm. He reported the largest turtle on record as being 188 mm Jong. Conant (1975) stated that C. p. marginata usually range in length from 115 to 140, the record length being 187 mm. Thus, the north- central Nogies Creek population falls within the range of carapace lengths that would be ex- pected of C. p. marginata. Figure 4 reveals a difference in the distribution of turtle sizes between the male and female subsamples. This difference is worth noting, but the growth relationships and other factors which may have created it are beyond the scope of the present study. Spring Activities That turtles respond predictably to tempera- ture has been indicated by several researchers. Sexton (1959) found that mass emigration of C. p. marginata would be initiated only when the temperature of the inlet was 8°C or higher. Ernst (1972) established the critical maximum temper- atures, and the temperature below which dor- mancy occurred. The number of turtles caught per unit of effort in Nogies Creek oscillated with major temperature changes. Greater catches corresponded with higher temperatures but there was often a lag period between the peaks in temperature and activity. Some authors, how- ever, have noted an acute ability to sense light. 56 THE CANADIAN FIELD-NATURALIST Noble and Breslau (1938) found hatchlings were attracted to high illumination. Ortleb and Sexton (1964) discovered that C. picta not only responded to light but were able to discriminate between light intensities of 0.1 foot candles. Light and temperature in the natural setting are usually closely linked and their respective effects may be confused. Legler (1954) reported that C. p. be//ii usually nested between 1700 and 1800 hours. Mahmoud (1968) observed nesting between 0500 and 0900 hours and later between 1600 and 2300 hours. The female C. p. marginata at Nogies Creek commenced daily nesting activities probably no earlier than 1400 hours and ceased leaving the water by about 1800 hours. This did not vary noticeably with air temperature. The annual initiation of nesting was also precise. In 1975 it was only 6 days later than in 1974. But 1975 had a remarkably warmer spring and if temperature is instrumental in inducing nesting behavior there should have been an appropriate early start of nesting. The uniformity of timing in daily and yearly activity would perhaps be more correctly attrib- uted to light than temperature. Sensitivity to photoperiod would adequately explain the tim- ing accuracy noted in this study. It would provide a much more dependable diurnal and annual clue than temperature. By having a set, safe activity period the turtles would not be fooled by short-term temperature changes when a long-term endeavor such as a summer’s incubation is at stake. It is possible that un- favorable temperature may discourage a photo- periodically induced turtle if there is potential danger to the individual or activity. Nogies ’ Creek C. p. marginata were also observed emerging in abnormal abundance following an afternoon shower, possibly to take advantage of the better digging conditions. Other environ- mental factors may occasionally override the basic photoperiodic clue. The long distance travelled to an “ideal” nest site by some Nogies Creek turtles and the high incidence of turtles found along several exposed sand banks suggests a form of homing. Further- more, marking turtles at aquatic capture sites revealed females as static during this their anticipated active time of the year. The ratio of males to females taken in nets favored males. Vol. 91 From recapture information, males are clearly more active than the females. The absence of females from the records may be normal if many female turtles were on or near shore. But in their movements to and from nesting sites they should have turned up in the netting results. This can be explained if the females are able to migrate toa chosen nesting site and return to the site of normal activity. A random net distribution would likely be sensitive to a randomly wander- ing population of female turtles yet could con- ceivably be ineffective against a directed move- ment. Many authors have documented homing in Chrysemys picta: Cagle (1944), Williams (1952), Gould (1959), Ortleb and Sexton 1964), Emlen (1969), and Ernst (1970b). Emlen (1969) pre- sented a strong argument for visual recognition of local topographic landmarks in orientation of C. picta. He pointed out Casteel’s (1911) findings of good visual discrimination in the species as well as a notable long-term memory. The long-distance travel to appropriate and hidden nesting sites described in this paper may also indicate homing, perhaps as has been hypothesized for marine turtles in the form of a return to the place of birth (Carr 1972) or as a return to a previously discovered suitable nest- ing location. Acknowledgments This research study is the result of secondary activities in the Muskellunge Research Project at Nogies Creek Research Station near Bobcay- geon, Ontario. We are grateful to the Canadian National Sportsmen’s Show, the Ontario Ministry of Natural Resources, and Grant A- 1705 National Research Council of Canada for financial support. We also extend our appre- ciation to Fergus McNeil who assisted in collecting the turtles, to Sophie Poray-Swinarski for constructing the figures, to F. R. Cook for his encouragement, and to J. P. Bogart for his suggestions and later criticism of the manu- script. Literature Cited Babcock, H.L. 1919. The turtles of New England. Me- moirs of the Boston Society of Natural History 8: 323-431. Bishop, S.C. and F. J. W. Schmidt. 1931. The painted turtles of the genus Chrysemys. Field Museum of Natural 1977 History Publications, Zoological Series 18: 123-139. Bleakney, J.S. 1958. Postglacial dispersal of the turtle Chrysemys picta. Herpetologica 14(2): 101-104. Brown, D.M. 1972. Heat units for corn in Southern Ontario. Ontario Ministry Agriculture and Food Fact- sheet. 4 pp. Cagle, F. R. 1939. A system of marking turtles for future identification. Copeia 1939(3): 170-173. Cagle, F.R. 1944. Home range, homing behavior, and migration in turtles. Miscellaneous Publication of the Museum of Zoology, University of Michigan 61: 1-34. Carr, A. 1952. Handbook of turtles. Cornell University Press, Ithaca, New York. 542 pp. Carr, A. 1972. Great reptiles, great enigmas. Audubon 74(2): 24-25. Casteel, D.B. 1911. The discriminative ability of the painted turtle. Journal of Animal Behavior 1(1): 1-28. Conant, R. 1975. A field guide to reptiles and amphibians. Houghton Mifflin Company, Boston, Massachusetts. - 429 pp. Crossman, E. J. 1956. Growth, mortality and movements of a sanctuary population of maskinonge (Esox mas- quinongy Mitchill). Journal of the Fisheries Research Board of Canada 13(5): 599-612. Emlen, S. T. 1969. Homing ability and orientation in the painted turtle Chrysemys picta marginata. Behavior 33: 58-76. Ernst, C.H. 1970a. The status of the painted turtle, Chrysemys picta, in Tennessee and Kentucky. Journal of Herpetology 4(1-2): 39-45. Ernst, C. H. 1970b. Homing ability in the painted turtle, Chrysemys picta (Schneider). Herpetologica 26(4): 399-403. Ernst, C. H. 1971a. Growth of the painted turtle, Chrys- emys picta, in southeastern Pennsylvania. Herpetologica 27(2): 135-141. Ernst, C. H. 1971b. Observations of the painted turtle, Chrysemys picta. Journal of Herpetology 5(3-4): 216-220. Ernst, C.H. 1971c. Chrysemys picta. Catalogue of the American amphibians and reptiles. Society for the Study of Amphibians and Reptiles: 106.1-106.4. Ernst, C. H. 1972. Temperature-activity relationships in the painted turtle Chrysemys picta. Copeia 1972(2): 217-222. Ernst, C.H. and R.W. Barbour. 1972. Turtles of the WHILLANS and CROSSMAN: MIDLAND PAINTED TURTLE, CENTRAL ONTARIO 57 United States. University Press of Kentucky, Lexington, Kentucky. 347 pp. Ernst, C. H. and E. M. Ernst. 1971. The taxonomic status and zoogeography of the painted turtle, Chrysemys picta, in Pennsylvania. Herpetologica 27(4): 390-396. Gould, E. 1959. Studies on the orientation of turtles. Copeia 1959 (2): 174-176. Hartman, W.R. 1958. Intergradation between two sub- species of painted turtles, genus Chrysemys. Copeia 1958(4): 261-265. Hartweg, N. 1944. Spring emergence of painted turtle hatchlings. Copeia 1944(1): 20-22. Legler, J. M. 1954. Nesting habits of the western painted turtle, Chrysemys picta bellii (Gray). Herpetologica 10(3): 137-144. Mahmoud, I. Y. 1968. Nesting behavior in the western painted turtle, Chrysemys picta bellii. Herpetologica 24(6): 158-162. Masat, R. J. and X. J. Musacchia. 1965. Serum protein concentration changes in the turtle, Chrysemys picta. Comparative Biochemical Physiology 16(2): 215-225. McCombie, A.M. 1959. Some relations between air temperature and the surface water temperatures of lakes. Limnology and Oceanography 4(3): 252-258. Muir, B.S. 1963. Vital statistics of Esox masquinongy in Nogies Creek, Ontario. I. Tag loss, mortality due to tagging, and the estimate of exploitation. Journal of the Fisheries Research Board of Canada 20(5): 1213-1230. Noble, G. K.and A. M. Breslau. 1938. The senses involved in the migration of young freshwater turtles after hatching. Journal of Comparative Psychology 25(1): 175-193. Ortleb, E. P. and O. J. Sexton. 1964. Orientation of the painted turtle, Chrysemys picta. American Midland Naturalist 71(2): 320-334. Pough, F.H. and M.B. Pough. 1968. The systematic status of painted turtles Chrysemys in the northeastern United States. Copeia 1968(3): 612-618. Sexton, O. J. 1959. Spatial and temporal movements of the painted turtle, Chrysemys picta marginata (Agassiz). Ecological Monographs 29(2): 113-140. Williams, J. E. 1952. Homing behavior of the painted turtle and musk turtle in a lake. Copeia 1952(2): 76-82. Received 23 June 1976 Accepted 21 November 1976 Germination Requirements of Alaskan Rosa acicularis R. DENSMORE and J. C. ZASADA Institute of Northern Forestry, USDA Forest Service, Fairbanks, Alaska Densmore, R.and J.C. Zasada. 1977. Germination requirements of Alaskan Rosa acicularis. Canadian Field-Naturalist 91(1): 58-62. Abstract. Germination requirements of Alaskan seeds of Rosa acicularis, a common shrub in the boreal zone of Asia and North America, were determined from laboratory experiments and observations of germination under outdoor conditions. Germination was rapid and complete at temperatures of 5°C to 20°C after 2 months of warm stratification and 3 months of cold stratification. Cold stratification alone or with a pretreatment of concentrated H,SO,, resulted in slow and incomplete germination. Laboratory and outdoor experiments indicated that most R. acicularis seeds take 2 years to germinate. Seeds develop and mature the first growing season, warm stratify the next growing season, cold stratify the following winter, and germinate in the spring shortly after snowmelt. Suggestions are made as to the overall reproductive strategy of R. acicularis and the role of its germination requirements. Rosa acicularis Lindl. (prickly rose) occurson and snowshoe hares. Fruits are also eaten by a broad range of sites in the boreal zone of Asia humans and are an excellent source of vitamin and North America (Figure 1). In Alaska this C. Rosaacicularis has potential as a revegetation species is commoninforestsandisabundantasa species for disturbed areas where food for successional species in disturbed areas. The wildlife and aesthetics are important considera- plant is an important food source for many tions. Little information is available in the animals, including microtine rodents, grouse, literature on regeneration from seed in this species; however, Babb (1959) recommended soaking seeds of R. acicularis in concentrated H,SO, for 1 h, then cold-stratifying at 6.5°C for 3 months, but he did not report what work was done as the basis for his recommendations. Other species of the genus Rosa that have been studied appear to have the same type of dormancy and similar germination require- ments, although depth of dormancy varies. Temperature-zone Rosa species required cold stratification to break dormancy and germ- inated poorly or not at all when kept at warm temperatures (Blundell and Jackson 1971; Crocker and Barton 1931; Nyholm 1955; Semeniuk and Stewart 1966; Svejda 1968). Pre- treatments of soaking in concentrated H,SO, (Blundell and Jackson 1971; Svejda 1968; United States Forest Service 1948) or warm stratification followed by cold stratification (Bouillene-Comhaire 1970; Rowley 1956; Sem- eniuk and Stewart 1966; Svejda 1968) often gave higher and more rapid germination than cold stratification alone. This paper describes the germination requirements of R. acicularis as determined from laboratory experiments and FIGURE |. Distribution of Rosa acicularis Lindl. is outlined 3 : : : on the map. Seeds for this study were collected near 4 Observations of germination of seeds subjected Fairbanks, Alaska. to outdoor conditions. 58 1977 Materials and Methods Hips! were collected in September 1972, May 1973, September 1974, and September 1975 near Fairbanks, Alaska (64°51’N, 147°44’ W). Achenes were separated from the pulp, washed, dried at room temperature for 48 h, and stored at 2-3°C in plastic bags. Laboratory germination experiments were conducted in controlled-temperature (+1°C) growth chambers with light/dark period of 16/8 h under white fluorescent tubes, light in- tensity 200 to 500 foot candles. Distilled water was used for all laboratory experiments. Seeds were germinated on vermiculite in plastic con- tainers with transparent lids and three or four 50-seed replications were used for each treat- ment. In laboratory tests, emergence of the radicle was regarded as germination, but seed- lings were transplanted to soil to check for normal development. To determine the conditions necessary to break dormancy, the following laboratory ex- periments were conducted on seeds collected in September 1974. (1) Seeds were cold stratified at 5°C for | year. Germination occurred at stratification tempera- tures. (2) Seeds were soaked for | h in concentrated H,SO,, rinsed, and stratified at S°C. The acid treatment reduced the thickness of the pericarp by half. On a portion of the seeds the pericarp was entirely removed in spots, and insome seeds the testa was also removed in these spots. Germination occurred at stratification temper- atures. (3) Seeds were warm stratified at 25°C for 115 days on cellulose pads and then placed on moist vermiculite at 5°C. Two weeks after germina- tion began at S°C, sets of replications were moved to 20°C and 10°/20°C?. One set of replications was retained at 5°C. In all experi- ments, seeds that had not germinated at the end 'The rose hip is an aggregate fruit enclosing several achenes. An achene is a one-sided fruit with the seed enclosed in a hard pericarp. This dispersal unit will hereafter be referred to as a seed. 2Diurnal alternation with a maximum of 20°C during the day for several hours, followed by a slow decline to a minimum of 10°C at night for several hours, after which temperatures slowly increased to 20°C. DENSMORE and ZASADA: ALASKAN ROSA ACICULARIS 59 of the experiments were dissected to determine the number of filled seeds. Germination per- centages are based on the number of filled seeds. To examine the response of the seeds to outdoor conditions, and to test planting methods for artificial revegetation, the following experiments were conducted. (1) Seeds collected in May 1973, from hips which had overwintered attached to the plant, were placed on soil in six pots, 25 seeds per pot, and covered with 2 cm humus and litter. Pots were placed outdoors in May. Seeds in the pots overwintered under the snow until 22 January. Then, because of time restrictions on the experi- ment they were dug out of the snow, brought indoors, and placed at 5°C to determine the response of the seed lot to the environmental conditions of the summer, fall, and one-half of the winter. (2) An attempt was made to shorten the normal 2-year period between planting and germina- tion. Seeds collected in early September 1974 were warm stratified for I’ month at 25°C and then planted outdoors on a mineral soil seedbed in early October. (3) To determine if any seeds would germinate the first spring after they were produced, after being subjected to a fall and winter of cold stratification but no warm temperatures, seeds were collected in September and planted out- doors on a mineral seedbed. Results and Discussions Seeds which were cold stratified at S5°C (experiment 1) germinated slowly and incom- pletely (Figure 2). Germination began at 5°C after 110 days. Seeds retained at 5°C continued to germinate, and germination rate of these seeds increased after 220 days, but after | year only 57% of the seeds had germinated. The germina- tion rate and capacity of seeds pretreated with acid (experiment 2) was very similar (Figure 2). The only effect of the acid treatment was that after 56 days, 4% of the seeds had germinated. These were seeds in which the acid had com- pletely removed sections of the seed covers, including the testa. Seeds which were warm stratified at 25°C for 118 days before being cold stratified (experiment 3) gave the most rapid and complete germination 60 THE CANADIAN FIELD-NATURALIST 70 ® Cold stratification 60 © Cold stratification with H,SO, pretreatment GERMINATION (Percent) 160 Vol. 91 180 200 220 240 260 280 300 320 340 360 DAYS FIGURE 2. Effect of cold stratification at 5°C and cold stratification with concentrated H,SO, pretreatment on germination of Rosa acicularis. Day zero is the day seeds were placed at 5°C, and germination occurred at stratification temperatures. (Figure 3). Germination began after 92 days at 5°C. Germination had reached 60% at 5°C after 107 days, when sets of replications were moved to 20°C and 10°/20°C. There were no signifi- cant differences in total germination between these seeds and the set of replications kept at 5°C. Apparently in seeds which are pretreated with warm stratification, metabolic processes involved in breaking of dormancy under cold stratification proceed at a relatively similar rate in the entire seed lot. Seeds then germinate quickly over a wide range of temperatures. Investigators (Blundell and Jackson 1971; Svejda 1968) working with other species of Rosa have reported that the role of acid or warm stratification pretreatments was to break down the pericarp, reducing mechanical resistance to embryo growth, or reducing the amount of inhibitors present in the pericarp. In R. acicul- aris the pericarp does not inhibit imbibition of moisture, and it apparently offers little resis- tance to embryo growth. After 2 or 3 months of cold or warm stratification, the seeds can be easily opened along the suture. In many seeds, the pericarp is split open and may fall off, but the seed remains dormant. The breakdown of the pericarp to reduce the amount of inhibitor present is probably not a major role of warm 100 } ® Germinated at 5°C O Moved to 20°C §0- © Moved to 10°/20°C ,-0-9~° 2 an 80 we a 4 a = 0 y; Cc a : 4 a 60 a Zz O50 0.05). Certain significant differences (F = 17.78; df= 4, 299; P<0.001) do occur among the populations in the pooled average number of anal fin rays with a low value of 12.42 for fish from Layton’s Lake and a high value of 14.25 for speci- mens from Large Lake (Table 3). The populations of N. crysoleucas from Layton’s Lake and Wood’s Pond are not significantly different (¢ = 1.33; P< 0.10) in anal fin ray number, but each had a count significantly lower than those of the other three populations. Large Lake and Long Lake, the most closely associated lakes of the series, supported populations that do not differ (P< 0.75) in number of anal fin rays. A significant difference does occur, however, between the population in Morice Lake and the other four populations. Discussion Notemigonus crysoleucas is known to have marked geographical variation in number of anal fin rays although other meristic series appear to be more constant (Hubbs 1921; Scott and Crossman 1973). Consequently, it is not unexpected that the number of dorsal and pelvic rays and the number of scale rows above the lateral line do not vary among the five populations examined, even though their habitats vary in certain limnological characteristics. The number of lateral line scales of the populations of Morice Lake, Long Lake, Large Lake, and Wood’s Pond also do not show significant differences and the pooled mean number (46.9) and range (42-49) agree well with the mean (47) and range (42-53) given by N Mean SE 14 15 15 = 75 12.96 0.08 21 2 72 13.17 0.09 14 = 40 13.25 0.10 2 = 24 12.63 0.15 2 = 93 12.42 0.07 Hubbs (1921) for specimens covering the range of the species. The population in Layton’s Lake, the most alkaline and eutrophic lake in the series does, however, possess a significantly lower number of lateral line scales. The number of anal fin rays, the meristic character in N. crysoleucas most susceptible to environmental modification (Hubbs 1921) does show the most extensive variation within the five populations. Long Lake and Large Lake are in the same drainage system and, at the closest point of contact, are separated by only 200 m. Although these lakes differ somewhat in their limnological characteristics, the two populations do not have significant differences in the meristic series examined in this study. Populations that occurred in these lakes in the past would have been eradicated by the inundation of these lakes by salt water as a result of the Saxby Gale of 1869 (Clair and Paterson 1976). Re-colonization of the lakes from sections of the drainage system which escaped the salt- water intrusion could have been from a single population and the limited time and extent of isolation since recolonization could have prevented expression of genetically controlled meristic dif- ferences. Morice Lake, which is limnologically similar to both Long Lake and Large Lake but lies in a different drainage system, supports a population of N. crysoleucas that has an anal fin ray number that differs significantly from that found in the other four populations. The two most diverse lakes in this series, Wood’s Pond and Layton’s Lake, contain fish that do not differ in the number of anal fin rays, although the average number is, in both cases, significantly lower than that for the other three lakes. The correlation between May temperature and anal fin ray number as established by Schultz (1926) would suggest that the May temperature in Wood’s Pond and Layton’s Lake would have to be about 5.5°C lower than in the other lakes to produce an effect of the observed magnitude if temperature is the factor that brings about the reduced number of anal rays in 1977 these two populations. Available data fail to reveal any obvious differences in the temperatures of comparable water masses in any of the five lakes at any time during the ice-free period. Further, year-to- year fluctuations in water temperature in the region are greater than between lake differences (J. B. Livingstone, unpublished manuscript, Mount Allison University). The lack of any apparent correlation between number of anal fin rays in the five populations and either the temperature or the other limnological characteristics of the lakes suggest that the observed differences are most probably not the result of obvious environmental differences between the habi- tats. In addition, the absence of differences in anal ray numbers between the two year classes examined in detail would appear to preclude short-term environ- mental shifts during critical periods of development as being responsible for the observed population differ- ences, unless it is assumed that such shifts were consistent in timing and intensity for each lake over at least a two-year period. The available data appear to be best interpreted by postulating genetic differences between the populations. Acknowledgments We thank all those people who gave so generously of their time in the collection of the specimens and J. Kerekes of the Canadian Wildlife Service for making available unpublished limnological data for Large Lake and Long Lake. This study was supported by the National Research Council of Canada through Grant A-6299. NOTES V7 Literature Cited Barlow, G. W. 1961. Causes and significance of morph- ological variation in fishes. Systematic Zoology 10: 105-117. Clair, T. and C. G. Paterson. 1976. Effect of a salt water intrusion on a freshwater Chironomidae community: a paleolimnological study. Hydrobiologia 48: 131-135. Hubbs, C. L. 1921. Geographical variation of Notemi- gonus crysoleucas—an American minnow. Transactions of the Illinois State Academy of Sciences 10: 147-151. Hubbs, C. L. 1926. The structural consequences of modifi- cations of the developmental rates in fishes, considered in reference to certain problems of evolution. American Naturalist 60: 57-81. Hubbs, C. L. 1941. Increased number and delayed develop- ment of scales in abnormal suckers. Papers of the Michigan Academy of Science, Arts and Letters 26: 229-237. Lee, C. L. and W. D. Williams. 1970. Meristic differences between two conspecific fish populations in Australian salt lakes. Journal of Fish Biology 2: 55-56. Lindsey, C. C. 1958. Modifications of meristic character- istics by light duration in kokanee, Oncorhynchus nerka. Copeia 1958: 134-136. Schultz, L. P. 1926. Temperature-controlled variations in the golden shiner, Notemigonus crysoleucas. Papers of the Michigan Academy of Science, Arts and Letters 7: 417-432. Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin 184. 966 pp. Taning, A. V. 1952. Experimental study of meristic charac- ters in fishes. Biological Review (Cambridge) 27: 169-193. Received 13 April 1976 Accepted 8 August 1976 Unreliability of Strip Aerial Surveys for Estimating Numbers of Wolves on Western Queen Elizabeth Islands, Northwest Territories FRANK L. MILLER and RICHARD H. RUSSELL Canadian Wildlife Service, Western and Northern Region, 10025 Jasper Avenue, Edmonton, Alberta T5J 1S6 Abstract. Numbers of wolves (Canis Jupus arctos) were obtained by aerial survey and by ground observation on western Queen Elizabeth Islands, Northwest Territories. Six transect-strip “census” aerial surveys were flown in March-April and July-August of each year from March 1972 to August 1974. The estimates based on aerial surveys were usually misleading. The behavior of observed wolves and their associations with other animals or objects that helped attract the observers attention to the wolves greatly influenced the number of observations. These reported shortcomings of aerial surveys must be considered during any future attempts at determining numbers of wolves. The so-called Melville Island wolf (Canis lupus arctos Pocock, 1935) that occurs on the Queen Elizabeth Islands, Northwest Territories, is the Canadian High Arctic form of the gray wolf, Canis lupus. This subspecies has apparently recently in- vaded Banks Island and replaced the Banks Island tundra wolf, C. /. bernardi Anderson, 1943 (Mann- ing and Macpherson 1958). It is likely that C. /. arctos now occurs on most or all islands south of Melville Sound, although the Baffin Island tundra wolf C. /. 78 manningi Anderson, 1943, is stillapparently found on Baffin Island. The Canadian High Arctic form of Canis apparently radiated out from the refugium of Pearyland after the Wisconsin glaciation (Macpher- son 1963). Information on High Arctic wolves is fragmentary, but existing reports (Parry 1821; Belcher 1855; M’Dougall 1857; M’Clintock 1859; Bernier 1910; Stefansson 1921; MacDonald 1954) suggest that historically wolves have not occurred in high numbers on the western Queen Elizabeth Islands. More recent observations (Macpherson 1961; Tener 1963; Riewe 1975) further emphasize the low numbers of wolves even during periods of high prey populations. High Arctic wolves prey mainly on Peary caribou ( Rangifer tarandus pearyi) and muskoxen (Ovibos moschatus). The future of C. /. arctos is questionable in view of the recent (1973-74) crash of both prey species and increasing human contact on the western Queen Elizabeth Islands. Man’s activities in the High Arctic will most likely encourage a wider range of wildlife inventories by involved agencies. The most practical method of counting arctic ungulates and large carnivores over THE CANADIAN FIELD-NATURALIST Vol. 91 extensive areas Is by aerial survey. The “transect-strip census” method is the design most often used for such counts. We flew six transect-strip aerial surveys between March 1972 and August 1974 to estimate Peary caribou and muskox numbers. Analysis of wolf observations made by us during the surveys shows that attempts to determine wolf numbers by this technique gave disparate estimates. Therefore, we present our observations of wolves obtained from aerial surveys and from additional sightings by observers on the ground in order to draw attention to the variation and possible error that can occur even from intensive (25% coverage) aerial strip surveys. Survey Area The Queen Elizabeth Islands included in our surveys are listed in Table 1 by descending order of size. The islands surveyed lie between latitudes 74° and 78° North and longitudes 95° and 124° West. The survey area, except western Melville Island is low- lying and mainly below 150m elevation. Western Melville Island is mostly mountainous terrain with many sites from 300 m to 1000 m above sea-level. The TABLE 1—Sightings of wolves on western Queen Elizabeth Islands, Northwest Territories, between March 1972 and August 1974 Wolf sightings 1972 1973 1974 Islands Size of surveyed island (km?) winter summer winter summer winter summer Melville 42220 15* 0(15)* 9(1) 14(22) 12 (7) 0(26) Bathurst 16090 1 0 (8) 0 Prince Patrick 15830 1(5) 12 (3) 0(10) 0(25) Mackenzie King 5100 0 0 Borden 2790 0 Eglinton 1550 0 0 0 8 1 (4) 0 Lougheed 1300 0 0 Byam Martin 1160 2 0 0 0 0 Vanier 1130 0 0 Cameron 1060 0 0 Brock 790 0 Emerald 550 0 0 0 Alexander 490 0 0 Massey 440 0 0 Little Cornwallis 410 0 0 0 Helena 330 0 0 Edmund Walker 82 0 0 Marc 56 0 0 Fitzwilliam Owen i 34 0 0 Eight Bears 18 0 0 *Sightings not in parentheses were obtained from aerial surveys and those sightings in parentheses were made by observers engaged in other activities. No entries (blank spaces) indicate that the islands were not surveyed during that period. 1977 terrain is open and for the most part is suitable for aerial surveys. Locally, broken terrain could hinder observation. Methods The islands were surveyed by use of a standard transect survey (Miller and Russell 1974). Surveys were flown on 13 days between 20 March and 7 April 1972, 8 days between 7 August and 24 August 1972, 18 days between 19 March and 15 April 1973, 18 days between 5 July and 21 August 1973, 12 days between 25 March and 17 April 1974, and 17 days between 18 July and 26 August 1974. Parallel flight lines were drawn on 1:250 000 scale topographical maps. In 1972 flight paths were at 6.4-km intervals. In 1973 and 1974 flight paths were also at 6.4-km intervals, except on Mackenzie King, Borden, and Brock islands where they were 12.8 km apart, and on Eglinton and Byam Martin islands the spacing was 3.2 km. The flight lines on Melville Island were oriented either east- west or north-south in each stratum to provide maximum contact with the coast for accurate naviga- tion. Flight lines were oriented east-west on all other islands although on Byam Martin and Eglinton islands, north-south surveys were added to provide double coverage in March-April 1973, then changed to all east-west lines for remaining surveys. A Helio Courier fixed-wing aircraft was used for all surveys, except in August 1972 when a Bell 206 turbo- helicopter was used. A 1.6-km strip, 0.8 km on each side of the aircraft, was surveyed. The 0.8-km strips were divided into two 0.4-km strips to determine the efficiency of observing within the 1.6-km strip. To mark the boundaries of each strip, wires were strung from an eye-bolt on the NOTES 719 wing to one on the fuselage of the Helio Courier. Lines marked on each observer’s window were aligned with corresponding tabs on the wires. At an altitude 150 m above ground, these tabs were checked against fuel drums located at 0.4-.and 0.8-km intervals from a reference point on the ground. Allowance was made for the blind spot beneath the aircraft so that an entire 0.8-km strip was visible on each side of the aircraft. Wildlife sightings were recorded as being within the two 0.4-km strips closest to the aircraft, within either of the two 0.4-km strips farthest from the aircraft, or outside both sets of strips (off transect). All survey flights were flown about 150 m above ground level according to altimeter readings except on western Melville Island where broken terrain forced us higher. Speeds ranged from 110 to 190 km/h, depending on the number of animals encountered. Observations were located on the survey maps and recorded on tape. At the end of each day the sightings were transcribed and located on a second map. Results and Discussion The distributions of wolves observed during our six aerial surveys and by observers on the ground are given in Table |. Table 2 gives numbers of wolves observed on each aerial survey, and estimated densities and numbers of wolves for all the islands and includes sizes of areas surveyed and distances flown. Observations in Table | show that during periods when no wolves were seen by aerial survey, wolves were seen by ground observers. Such discrepancies between aerial and ground observations further support the apparent observational error resulting from aerial survey of wolves. Tener (1963) saw 18 wolves during his aerial survey of the entire Queen TABLE 2—Observed and estimated numbers of wolves on western Queen Elizabeth Islands, Northwest Territories, obtained from six aerial surveys Estimates Observed density* by strip location of total numbers (wolves/ 1000 km?) based on Area Distance Total widths of strips surveyed flown wolves Outer strips Inner strips All strips Survey period (km?) (km) seen (total0.8km) (total0.8km) (total 1.6 km) 0.8km_ 1.6km Mar.-Apr. 1972 44930 7020 17 0.0 3.0 1.5 135 67 Aug. 1972 26240 4100 0 0.0 0.0 0.0 0 0 Mar.-Apr. 1973 91430 13930 11 1.0 0.0 0.5 0 46 July—Aug. 1973 61310 10000 25** 3.2 0.0 1.6 0 98 July-Aug. 1973 61310 10000 34*** 4.2 0.0 2.1 0 129 Mar.-Apr. 1974 50400 7920 14 DED, 0.0 1.1 0 55 July-Aug. 1974 67800 10340 0 0.0 0.0 0.0 0 0 *Density was determined by multiplying distance flown (km) by width of transect strip (km), which equals area censused (km?). then total wolves seen on transect was divided by the area censused: e.g.. 7020 X 0.8 = 5616 km’, then 17/5616 = 0.0003 wolves/km* or 3.0 wolves/ 1000 km’. **Observations do not include newborn pups. ***T otal wolves seen includes nine newborn pups. 80 THE CANADIAN FIELD-NATURALIST Elizabeth Group in summer 1961 and only three of those wolves were on western Queen Elizabeth Islands. Coverage was only 6% for the 1961 survey. At that time on the western islands Peary caribou were estimated to number 24 320, about 10 times as numerous as we estimated in 1974 (2676). Muskoxen occurred at similarly estimated numbers (2161 in 1961 and 2704 in 1974). Of the 202 total sightings of wolves made both from the air and from the ground between March 1972 and August 1974 (Table 1), 69% (140) wolves were seen in 24 packs. Pack size averaged 5.8 and ranged from 2 to 15. Twenty-four percent (49) of the wolves seen were in six family groups of adults and pups, five groups in summer 1973 and only one in summer 1974. Family groups averaged 4.2 adults/4.0 pups, and adults/ pups varied as follows: 2/5, 3/2, 4/2, 5/1, 5/10, and 6/4. Two dens were found on Melville Island and one on Prince Patrick Island in 1973. Only the den on Prince Patrick Island was occupied in summer 1974. Only 6% (13) of the wolves seen were solitary. Of all the wolves seen (Table 1) 14% were stalking muskoxen, 12% were feeding on muskox carcasses, 9% were stalking caribou, 17% were at wolf dens, 17% were travelling and not associated with anything that would have helped to attract our attention to them, and 31% were seen at permanent or temporary camp sites. Only 76 wolves were seen on 53 310 km of flight paths during the six aerial surveys (Tables | and 2): 53% were in packs, 35% were in family groups (20% at den sites), and 12% were solitary wolves. Of the 76 wolves seen, 28% (21) were stalking muskoxen, 22% (17) were feeding on muskox carcasses, 3% (2) were stalking caribou, 35% (27) were in family groups (15 at den sites and 12 travelling cross-country), and 12% (9) were travelling and not associated with any other animals or objects. The variations in the estimates (Table 2) established the inaccuracy of the estimates, but we are unable to calculate correction factors as we cannot verify the accuracy of any of the estimates. Our inability to determine how many of the wolves sighted by ground observers represented different individuals prohibits the establishment of a ratio of wolves seen from the ground to wolves seen from the air. Sucha ratio would have allowed evaluation of errors in the estimates obtained from aerial surveys. It is clear, however, that transect-strip aerial surveys are not reliable for determining wolf populations on the open tundra of the High Arctic. Some of the problems inherent in aerial surveys of large mammals (Graham and Bell 1969) particularly apply to our surveys of wolves. The size and relative lack of color contrast between pelage and back- ground, especially in winter, reduce the visibility of wolves from the air. In addition the behavior of the Vol. 91 wolves can influence their chances of being seen from the air. Although we do not have quantitative measurements, ground observations of wolves reveal that wolves will sometimes remain stationary during overflights by aircraft. The effect that arbitary selec- tion of transect strip widths has on subsequent estimation is shown in the estimates by strip width (Table 2). We cannot explain the unexpected pattern of occurrence of wolves on the outer and inner strips (Table 2). Our results do not follow the expected pattern of most observations of inconspicious animals on narrow transects (Pennycuick 1969). We suggest that some wolves had moved from the inner to the outer strips before being sighted and/or remained stationary on the inner strips and were not sighted. These conditions and the above factors, and possibly many more, contribute to confounding the observations of wolves by aerial survey. Our consistently low counts of wolves do suggest that wolves are in low numbers throughout the western Queen Elizabeth Islands. The current num- bers of Peary caribou and muskoxen could not sustain high numbers of wolves, and alternate food sources are often scarce. It is our opinion, however, that even the number of wolves seen on each aerial survey is not necessarily representative of the true number of wolves present. Therefore, resultant estimates would often be erroneous; this condition must be borne in mind by future observers. Acknowledgments For logistic help we thank Atmospheric Environ- mental Service, Department of the Environment; National Museum of Natural Sciences, High Arctic Research Station; Panarctic Oils Limited; and Polar Continental Shelf Project, Department of Energy, Mines and Resources. We thank other observers and members of Canadian Wildlife Service field parties that helped us obtain this information: G. A. Calder- wood, M. V. Channing, P. L. Madore, J. W. Max- well, G. R. Parker, L. S. Prevett, H. J. Russell, G. D. Tessier, D.C. Thomas, and D.R. Urquhart. A. Gunn, Canadian Wildlife Service, critically read the manuscript. Literature Cited Belcher, E. 1855. The last of the Arctic voyages: being a narrative of the expedition in HMS Assistance under the command of Capt. Sir Edward Belcher, C.B., in search of Sir John Franklin, during the years 1852, 1853, and 1854. With notes on the natural history by Sir John Richardson, Prof. Owen, Thomas Bell, J. W. Slater, and Lovell Reeve. L. Reeve, London. 383 pp. Bernier, J. E. 1910. Report on the Dominion Government Expedition to the Arctic Islands and Hudson Strait on board the C.G.S. Arctic in 1908-1909. Government Loy Printing Bureau, Ottawa. 529 pp. Graham, A. and R. Bell. 1969. Factors influencing the countability of animals. Jn Proceedings of the workshop on the use of light aircraft in wildlife management in East Africa. Edited by W.G. Swank, R. M. Watson, G. H. Freeman, and T. Jones. Special Issue East African Agri- cultural and Forestry Journal. 108 pp. Macdonald, S. D. 1954. Report on biological investiga- tions at Mould Bay, Prince Patrick Is., N.W.T., in 1952. Annual Report of the National Museum of Canada, 1952-53, Bulletin 132: 214-238. Macpherson, A. H. 1961. On the abundance and distri- bution of certain mammals in the Western Canadian Arctic Islands in 1958-9. Arctic Circular 14(1): 1-16. Macpherson, A. H. 1963. The origin of diversity in mam- mals of the Canadian Arctic tundra. Systematic Zoology 14(3): 153-173. Manning, T. H. and A. H. Macpherson. 1958. The mam- mals of Banks Island. Arctic Institute of North America Technical Paper Number 2: 1-74. M’Clintock, F. L. 1859. The voyage of the Fox in the Arctic Seas: A narrative of the discovery of the fate of Sir John Franklin and his companions. J. Murray, London. 403 pp. M’Dougall, G. F. 1857. The eventful voyage of H. M. Discovery ship Resolute to the Arctic regions in search of Sir John Franklin and the missing crews of H. M. Discovery ships Erebus and Terror 1852, 1853, 1854. NOTES 81 Longmans, London. 529 pp. Miller, F. L. and R.H. Russell. 1974. Aerial surveys of Peary caribou and muskoxen on western Queen Elizabeth Islands, Northwest Territories, 1973. Canadian Wildlife Service Progress Note Number 40: 1-18. Parry, W. E. 1821. Journal of a voyage for the discovery of a Northwest passage from the Atlantic to the Pacific: per- formed in the years 1819-20, in His Majesty’s ships Hecla and Griper, under the Orders of William Edward Parry... with an appendix, containing Scientific and Other observations. John Murray, London. 309 pp. Pennycuick, C. J. 1969. Methods of using light aircraft in wildlife biology. Jn Proceedings of the workshop on the use of light aircraft in wildlife management in East Africa. Edited by G. W. Swank, R. M. Watson, G. H. Freeman, and T. Jones. Special Issue East African Agricultural and Forestry Journal. 108 pp. Riewe, R.R. 1975. The High Arctic wolf in the Jones Sound region of the Canadian High Arctic. Arctic 28(3): 209-212. Stefansson, V. 1921. The friendly arctic. Macmillan Co., New York. 784 pp. Tener, J.S. 1963. Queen Elizabeth Islands game survey, 1961. Canadian Wildlife Service Occasional Paper Number 4: 1-50. Received 19 March 1976 Accepted 13 June 1976 The Flowering Phenology of Common Vascular Plants at Bailey Point, Melville Island, Northwest Territories GERALD R. PARKER Canadian Wildlife Service, Box 1590, Sackville, New Brunswick E0OA 3C0 A collection of the vascular flora at Bailey Point, Melville Island (74°58’ N, 115°01’ W) was made during the course of studies of caribou (Rangifer tarandus pearyi) and muskoxen (Ovibos moschatus) from 24 June to 20 August 1974. That collection is preserved in the Vascular Plant Herbarium of the Bio- systematics Research Institute, Ottawa. Cody et al. (1976) have reported on new additions to the vascular flora of Melville Island resulting from the collection. Throughout the period of observation, notes were kept on when the most common plants came into flower, the period of peak bloom, and when the last flowers of a species were seen. A search of the literature showed a paucity of information on the flowering phenology of vascular plants in the high Arctic. Existing records usually include the date of first flower for a few species only (Savile 1959, 1961, 1964; Parmelee 1963; Beschel 1963). Bruggemann and Calder (1953) provide a useful comparison of first- flowering dates for 16 vascular species at four locations in the Canadian Arctic. Records of flowering periods are useful in docu- menting species differences in phenology during a single growing season at specific locations. Broader regional differences in phenology may become apparent when more data become available. Such records are also useful to those persons planning to study certain species in specific northern locations. The period of flower, with approximate date of peak bloom, is shown in Figure | for the most common and conspicuous vascular plants at Bailey Point from 28 June to 15 August 1974. The first vascular plant in flower was Saxifraga oppositifolia; the last to appear was Senecio con- gestus. The period when the most species were in peak flower was 18 to 25 July. The degree of habitat 82 J SPECIES Saxifraga oppositifolia Ranunculus nivalis THE CANADIAN FIELD-NATURALIST Vol. 91 UNE JULY AUGUST 28 29 301234 56 7 8 910 111213 1415 16 1718 19 20 21 22 23 24 25 26 27 28 29 3031 123456 7 8 9 10 11213 1415 SS RS Salix arctica Parrya arctica Draba alpina Saxifraga flagellaris Eriophorum triste Potentilla hyparctica Alopecurus alpinus Luzula nivalis Pedicularis arctica Saxifraga caespitosa Petasites frigidus Cerastium arctium Eriophorum Scheuchzeri Cardamine bellidifolia Dryas integrifolia Saxifraga nivalis Oxyria digya Geum Rossii Papaver radicatum Cassiope tetragona Chrysoplenium tetrandrum Ranunculus Sabinei Eutrema Edwardsii Saxifraga Hirculus Luzula confusa Carex misandra Carex stans Caltha palustris Saxifraga tricuspidata Taraxacum pumilum Potentilla Vahliana Melandrium apetalum Taraxacum hyparcticum Saxifraga cernua Pedicularis sudética Cochlearia officinalis Astralagus alpinus Stellaria longipes Polygonum viviparum Pleuropogon Sabinei Arnica alpina Ranunculus hyperboreus Senecio congestus @ Approximate peak bloom (CR SSI ST ET SE FIGURE 1. Approximate period of flower and peak bloom for 45 common vascular plants at Bailey Point, Melville Island from 28 June to !5 August 1974. selectivity by a species often dictated the length of bloom. The flowers of Ranunculus nivalis may be seen shortly after those of Saxifraga oppositifolia on plants growing on exposed turfy tundra sites; they may also be among the last flowers to bloom on plants in adjacent late snowbeds. In contrast, the aquatic buttercup, Ranunculus hyperboreus, displayed an extremely short period of flower of only 5 days. The dates of first flower for representative vascular plants at Isachsen, Ellef Ringnes Island, in 1960 (Savile 1961) were considerably earlier than first- flowering dates for the same species at Bailey Point in 1974. Mean daily temperatures for June and July, however, are slightly lower at Isachsen than those at Mould Bay, reported as 0.0°C and 2.1°C, respec- 1977 tively, from 1951 to 1960 (Thompson 1967). Mould Bay is on Prince Patrick Island, approximately 150 km northwest of Bailey Point. Savile (1961) reported that 1960 was a typical growing season. He (Savile 1971) also suggested the better plant growth at Mould Bay may be explained by the greater number of sunshine hours there than at Isachsen. In 1974 mean temperatures for June and July at Mould Bay were -1.8°C and 2.0°C; this suggests that an abnormally cool month of June delayed plant development and subsequent flowering dates at Bailey Point by as much as 10 to 14 days. Literature Cited Beschel, R. E. 1963. Geobotanical studies on Axel Heiberg Island in 1962. Preliminary Report 1961-1962. By F. Muller. Jacobsen-McGill Arctic Research Expedition 1959-1962. Axel Heiberg Island Research Reports. McGill University, Montreal. pp. 119-215. Bruggemann, P.F. and J. A. Calder. 1953. Botanical in- vestigation in Northeast Ellesmere Island, 1951. Canadian Field-Naturalist 67(4): 157-174. Cody, W.J., G.R. Parker, and D.B.M. Lamperd. NOTES 83 1976. Additions to the vascular flora of Melville Island, Franklin District, Northwest Territories. Canadian Field- Naturalist 90(1): 70-72. Parmelee, J. A. 1963. Mycological studies in 1961. Pre- liminary Report 1961-1962. By F. Muller. Jacobsen- McGill Arctic Research Expedition 1959-1962. Axel Heiberg Island Research Report. McGill University, Montreal. pp. 173-181. Savile, D.B.O. 1959. The botany of Somerset Island, District of Franklin. Canadian Journal of Botany 37: 959-1002. Savile, D.B. O. 1961. The botany of the northwestern Queen Elizabeth Islands. Canadian Journal of Botany 39: 909-942. Savile, D. B. O. 1964. General ecology and vascular plants of the Hazen Camp area. Arctic 17(4): 237-256. Savile, D. B.O. 1971. Microclimate and plant growth at Isachsen and Mould Bay. Arctic 24(4): 306-307. Thompson, H. A. 1967. The climate of the Canadian Arctic. In The Canada Year Book, 1967. The Dominion Bureau of Statistics, Ottawa. pp. 55-74. Received 12 July 1976 Accepted 18 October 1976 Gestation, Litter Size, and Number of Litters of the Red Squirrel (Tamiasciurus hudsonicus) in Quebec JEAN FERRON! and JACQUES PRESCOTT? 1Département des sciences pures, Université du Québec, Rimouski, Québec GSL 3A1 2Jardin Zoologique de Québec, M.T.C.P., 8191 Avenue du Zoo, Orsainville, Québec GIG 4G4 In the course of ethological studies at the Station de Biologie de Université de Montréal, at St. Hippolyte, Quebec, from November 1971 to October 1974, information concerning the reproductive biology of Tamiasciurus hudsonicus in eastern Canada was gathered by the authors. Daily observation of captive squirrels (three males and three females) in a large outdoor enclosure (2.4 m high with a floor surface of 4.5 X 3.7 m) allowed one of us (J. F.) to record the exact day of copulation for one female, 10 April 1973 between 1300 and 1400 hours, and of parturition 35 days later on 15 May 1973 between 0000 and 0700 hours. That the female mated only on 10 April is in agreement with Smith (1968) who found that the oestrous female red squirrel is receptive for only | day. This is the first direct observation of the length of the gestation period for T. hudsonicus; it is shorter than the approximation of 40 days given by Hamilton (1939). Between June 1972 and August 1974 seven gravid female red squirrels were live-trapped at St. Hippo- lyte. They were placed in individual wire cages until parturition. They gave birth to 38 young (Table 1). The average litter size at birth was 5.4 (range 4 to 8, N=7); this is higher than 3.2 (N= 9, based on nest young) observed by Davis (1969), 3.9 (N = 82, based on placental scars) by Kemp and Keith (1970), 4.5 (N= 74, based on placental scars, embryos, nest young) by Layne (1954), 4.4(N = 83, based on corpora lutea counts) by Millar (1970), 3.3 (N = 24, based on placental scars, embryos, nest young) by Smith (1968), and 4.0 (N = 20, based on placental scars) by Wood (1967). From Table 1, a sex ratio at birth was calculated. We found a ratio of 1.37 males to 1.00 females (N = 38); this does not differ significantly from 1:1 ratio (x = 0.97, P > 0.05). Davis (1969) also found a sex ratio of 1:1 at birth for 7. hudsonicus in Saskatchewan. From year-round observation of females in the wild 84 THE CANADIAN FIELD-NATURALIST TABLE 1—Date of birth and litter composition of Tamia- sciurus hudsonicus at St. Hippolyte, Quebec. Young were counted and sexed shortly after parturition Date of birth Number of young 36 29 Total 2 June 1972 3 2 5 25 April 1973 3 2 5 28 April 1973 3 yp) 5 11 May 1974 2 2 4 2 June 1974 4 0 4 9 July 1974 6 1 7 31 July 1974 1 7 8 Total 22 16 38 and in a captive group kept under semi-natural conditions, it appears that females have only one litter per year. Wrigley (1969) suggests the existence of two breeding seasons in southern Quebec but did not have any observations of individuals having two litters per year. Layne (1954), however, found that in eastern United States red squirrels frequently produce two litters a year. In the northwestern part of North America, only one litter per year has been reported by Davis (1969), Dolbeer (1973), Hatt (1943), Kemp and Keith (1970), Smith (1968), Soper (1942), and Wood (1967). The short length of the mild season in northern regions and at high elevations probably limits production to one litter per year. This is supported by Millar (1970) who recorded two litters in British Columbia during a particularly mild year, and one litter during a normal colder year. We thank Professor Paul Pirlot for helpful com- ments, Jessica Pottier for reading the manuscript, and Vol. 91 the National Research Council of Canada and the Station de Biologie de Université de Montréal for financial assistance. Literature Cited Davis, D. W. 1969. The behavior and population dynamics of the red squirrel, Tamiasciurus hudsonicus, in Saskatch- ewan. Ph.D. thesis, University of Arkansas, Fayette- ville. 222 pp. Dolbeer, R. A. 1973. Reproduction in the red squirrel (Tamiasciurus hudsonicus) in Colorado. Journal of Mammalogy 54: 536-540. Hamilton, W. J., Jr. 1939. Observations on the life history of the red squirrel in New York. American Midland Naturalist 22: 732-745. Hatt, R. T. 1943. The pine squirrel in Colorado. Journal of Mammalogy 24: 311-345. Kemp, G. A. and L. B. Keith. 1970. Dynamics and regula- tion of red squirrel (Tamiasciurus hudsonicus) popula- tions. Ecology 51: 763-779. Layne, J. N. 1954. The biology of the red squirrel, Tamia- sciurus hudsonicus loquax (Bangs), in central New York. Ecological Monographs 24: 227-267. Millar, J.S. 1970. Variations in fecundity of the red squirrel, Tamiasciurus hudsonicus (Erxleben). Canadian Journal of Zoology 48: 1055-1058. Smith, C. C. 1968. The adaptative nature of social organi- zation in the genus of tree squirrels Tamiasciurus. Eco- logical Monograph 38: 31-63. Soper, J.D. 1942. Mammals of Wood Buffalo Park, northern Alberta and District of MacKenzie. Journal of Mammalogy 23: 119-145. Wood, T. J. 1967. Ecology and population dynamics of the red squirrel (Tamiasciurus hudsonicus) in Wood Buffalo National Park. M.Sc. thesis, University of Saskatchewan, Saskatoon. 97 pp. Wrigley, R. E. 1969. Ecological notes on the mammals of southern Quebec. Canadian Field-Naturalist 83: 201-211. Received 9 December 1975 Accepted 19 July 1976 The Fern Woodsia obtusa (Spreng.) Torrey in Ontario DONALD M. BRITTON Department of Botany and Genetics, University of Guelph The author, in the company of M. Coulthart, visited Frontenac Provincial Park north of Kingston on 30 July 1975, in an attempt to find Asplenium trichomanes L. growing together with A. platyneuron Oakes. It was a hot summer day and we noted that Cystopteris fragilis Bernh. growing in rocky crevices appeared completely brown and dead from the pro- longed summer drought. As we were returning to the , Guelph, Ontario NIG 2W1 car after walking along the shore of Black Lake immediately to the west of Devil’s Lake in Bedford Township, Frontenac County, we crossed a rough wall of granite boulders and a frond of a fern that was completely green caught my attention in amongst the dead fronds of C. fragilis. It looked like Woodsia obtusa, but this species is considered to be absent from Ontario (Lafontaine 1973); it was carefully put in a 1977 press for study back at Guelph. This specimen, Britton 3584 (OAC), indeed proved to be Woodsia obtusa (Spreng.) Torrey. The author returned to the location on 2 September 1975 with A.C. Jermy, British Museum (Natural History) in order to ascertain how many plants were present, decide if the species was native or introduced to the area, see if there were other colonies near by, and take photographs. We were able to count approx- imately 12 distinct clones growing in and under granite boulders and the colony covered a narrow band of about 46 m (50 yards) in length. The plants were in the semi-shade of Fraxinus americana, Carya ovata, Xanthoxylum americanum, and small Ulmus americana. The young trees appeared to be about 15-20 years old, and the Woodsia appeared to be well established under their canopy. Our impression was that the species was adventive to the area in the same sense that Asplenium platyneuron grows in disturbed or young second-growth sites, but at the same time the Woodsia is now well established, the colony being perhaps 10 years old. A specimen, Jermy 12424(BM), was collected. Woodsia obtusa is a very common species in north- eastern United States extending north to southern Maine, southern Quebec (Lafontaine 1973), New York, and Ohio. Brown (1964) commented that in a greenhouse it “grew like a weed and propagated itself over the benches, even growing in the cinders,” quite unlike the other species of Woodsia whichare difficult to cultivate. Wherry (1961) notes that it grows on various rocks, often invading masonry, the soil circumneutral to subacid. The species has been somewhat of a mystery plant in Ontario. There are two sheets in QK. One is from the herbarium of J. Macoun (Acc. No. 02736) and the other is labeled Ex. NOTES 85 Coll. Nicol (Acc. No. 2735), but neither sheet has any locality data. Naturalists have been puzzled by the pictures in Native ferns of eastern North America (Metcalfe 1963) with the caption “In Ontario the Obtuse Woodsia is much rarer than the rusty one.” The query has been raised as to whether this picture is really of Woodsia oregana var. cathcartiana, which 1s fairly abundant on Manitoulin Island, or did Metcalfe know of a colony of W. obtusa in Ontario? One might speculate that the range of Woodsia obtusa does expand into Ontario in periods of warmer winters and has done so in the past. It could be exhibiting the same characteristics as does another southern species, Asplenium platyneuron which seems to be occurring at more and more Ontario stations in recent years; itis now known from five sites in Norfolk County and has recently been collected in Elgin County, Brant County, and Waterloo County. In each case it appears to be a recent arrival. Literature Cited Brown, D. F. M. 1964. A monographic study of the fern genus Woodsia. Nova Hedwigia 16: 1-154. Lafontaine, J.D. 1973. Range extension of the Blunt-lobed Woodsia, Woodsia obtusa (Spreng.) Torr. (Poly- podiaceae), in Canada. Canadian Field-Naturalist 87(1): 56. Metcalfe, B. 1963. Section Two. /n Native ferns of eastern North America. 3rd edition (revised). The Canadian Audubon Society and the Federation of Ontario Natur- alists, Toronto and Don Mills, Ontario. 46 pp. Wherry, E.T. 1961. The fern guide. Doubleday and Company, Garden City, New York. Received 27 April 1976 Accepted 30 June 1976 First Report of the Tiger Trout Hybrid, Salmo trutta Linnaeus X Salvelinus fontinalis (Mitchill), in Alberta JAMES H. ALLAN Fish and Wildlife Division, Department of Recreation, Parks and Wildlife, Red Deer, Alberta T4N SY5 The tiger trout hybrid (Salmo trutta Lin- naeus X Salvelinus fontinalis (Mitchill) ) occurs rarely under natural conditions (Brown 1966). Although there is evidence of ecological and reproductive segre- gation of the brown trout (Sa/mo trutta) and the brook trout (Salvelinus fontinalis) there are areas in streams where the two species intermix, particularly during the coincidental spawning seasons in late October and early November (Marshall and Mac- Crimmon 1970; Nyman 1970; Vincent and Miller 1969). Although the opportunity for natural hybridi- zation may be fairly frequent, hybrids are rarely taken because of low fertility and survival of offspring (Buss and Wright 1956; Seguin 1957). The tiger trout hybrid 86 THE CANADIAN FIELD-NATURALIST Vol. 91 FiGuRE |. Tiger trout hybrid (Salmo trutta X Salvelinus fontinalis) taken from the North Raven River on 4 October 1974. has not previously been reported for Alberta (Paetz and Nelson 1970; Scott and Crossman 1973), though there are a number of streams where the brown trout and brook trout occur sympatrically. The North Raven River (52°10’ N, 114°41’ W) has the highest known populations in Alberta (i.e., in excess of 800 trout per kilometre in some sections, according to C. Shirvell (1972. Survey of Stauffer Creek and habitat program. Fish and Wildlife Division, Alberta Depart- ment of Lands and Forests Manuscript Report. 109 pp.)). Since 1972 over 11 000 specimens of both species have been examined during studies of this stream. No trout have been introduced to the North Raven River since 1950 and no hybridization experi- ments have taken place at the Raven Rearing Station (J. C. Barnhardt, personal communication). On 4 October 1974 a single specimen, presumed of the tiger trout hybrid, was taken from the North Raven River by electrofishing (Figure 1). The speci- men was 120 mm in fork length, weighed 23.2 g, and was 1+ years old. It was golden-brown with broad, dark brown vertical vermiculations on the lateral surfaces, which extended from the opercula to the caudal penduncle. The dorsal and adipose fins had irregular dark brown spots. Some meristics for the specimen are as follows: teeth on both the head and shaft of the vomer; gill rakers 15; branchiostegal rays 9; pelvic fin rays 9; pectoral fin rays 12; main dorsal fin rays 11; scales one row above lateral line 162; scales above lateral line to posterior insertion of dorsal fin Sl The specimen’s characteristics correspond to the photographs and description of the tiger trout hybrid given by Buss and Wright (1956, 1958) and Brown (1966). A.C. Sinclair, Superintendent of the Sam Livingston Fish Hatchery, Calgary, Alberta and J. S. Nelson, University of Alberta, Edmonton, Alberta confirmed the identification. The specimen is in the fish collection at the Sam Livingston Hatchery. Literature Cited Brown, C. J.D. 1966. Natural hybrids of Salmo trutta X Salvelinus fontinalis. Copeia 3: 600-601. Buss, K. and J. E. Wright, Jr. 1956. Results of species hybridization within the family salmenidae. Progressive Fish-Culturist 18(4): 149-158. Buss, K. and J. E. Wright, Jr. 1958. Appearance and fertility of trout hybrids. Transactions of the American Fisheries Society 87: 172-181. Marshall, T. Lawrence and Hugh R. MacCrimmon. 1970. Exploitation of self sustaining Ontario stream populations of brown trout (Sa/mo trutta) and brook trout (Sa/velinus fontinalis). Journal of the Fisheries Research Board of Canada 27(6): 1087-1102. Nyman, O. L. 1970. Ecological interaction of brown trout, Salmo trutta L. and brook trout, Sa/velinus fontinalis (Mitchill) in a stream. Canadian Field-Naturalist 87(4): 343-350. Paetz, M. J. and J.S. Nelson. 1970. The fishes of Alberta. Department of Mines and Minerals, Edmonton, Alberta. 282 pp. Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin 184. 966 pp. Séguin, Louis-Roche. 1957. Scientific fish culture in Quebec since 1945. Transactions of the American Fisher- ies Society 18(4): 136-143. Vincent, Robert E. and William H. Miller. 1969. Altitud- inal distribution of brown trout and other fishes in a head- water tributary of the South Platte River, Colorado. Ecology 50(3): 464-466. Received 24 March 1976 Accepted 13 August 1976 1977 NOTES 87 The Cattle Egret in British Columbia R. WAYNE CAMPBELL! and WAYNE C. WEBER? British Columbia Provincial Museum, Victoria, British Columbia V8W IAI 2Department of Zoology, Mississippi State University, Mississippi State, Mississippi 39762 The recent range expansion of the Cattle Egret (Bubulcus ibis) in North America has aroused wide interest. Since its arrival in Florida in the early 1940s (Palmer 1962), the species has been recorded in nearly all American states and Canadian provinces, and is a well established breeder in the southeastern United States (Rice 1956; Davis 1960; Blake 1961; Crosby 1972); it has also bred locally west of the Rockies, in southern California (Small 1974), and New Mexico (Witzeman et al. 1975). Cattle Egrets have now extended their range in western North America into British Columbia. This note reports 17 sightings for the province, involving at least 11 or 12 different birds; two of these are supported by specimens and another four by photographs (see Campbell and Stirling 1971). Table 1 lists all known records of Cattle Egrets for British Columbia. Details of all records are on file at the British Columbia Provincial Museum. The 1970 reports are questionable since two observations remain unconfirmed, and for the third (12 December) there was some uncertainty over the identification. The Lower Mainland records therefore were not listed by Campbell et al. (1972) although detailed field notes by the Webers strongly suggest a Cattle Egret. The first positive record for the province was on 19 November 1973 when the Goodwills obtained a roll of 16-mm color movie film of a Cattle Egret probing for food in manure in a pasture field with sheep and cattle. All records in 1973 and 1975 were from the vicinity of Vancouver Island, whereas 1974 reports were from widely separated localities. Egrets most often asso- ciated with domestic stock (cattle, sheep, horses, and even chickens) in farming areas (e.g., Saseenos, Oyster TABLE 1—Records of Cattle Egrets in British Columbia. sVI = southern Vancouver Island; LM = Lower Mainland (Vancouver area); wcVI = west coast Vancouver Island; scINT = south-central Interior British Columbia; ecVI = east coast Vancouver Island; BCPM = British Columbia Provincial Museum Observer Documentation Date Locality Area 26 Nov. 1970 West of Victoria sVI 12 Dec. 1970 Iona Island LM 21 Dec. 1970 Lulu Island LM 15-19 Nov. 1973 Saseenos sVI 8 Dec. 1973 Tofino weVI 15 Dec. 1973 Stubbs Island weVI 19 Nov. 1974 Lake Cowichan sVI 21 Nov. 1974 Pachena Point weVI 22 Nov. 1974 Salmon Arm scINT 16-31 Dec. 1974 Pitt Meadows LM 21 Dec. 1974 Burnaby Lake LM 22 Dec. 1974 Mud Bay LM 7 Jan. 1975 Denman Island ecVI 23-26 Nov. 1975 Saseenos sVI 24-29 Nov. 1975 Sooke sVI 27 Nov.-6 Dec. 1975 Oyster River ecVI 8-14 Dec. 1975 Nanaimo ecVI fide W. J. Schick W.C. Weber; R. R. Weber B. Gladstone A. Wickheim; J. E. V. Goodwill, M. Goodwill fide A. Dorst fide A. Dorst A. Allan F. G. Hart; R. Noble W.D. Munro B. vanDrimmelin; D. McDermott; S. H. Robinson, W. C. Weber; D. M. Mark S. Mooney; B. P. Watts H. Lamb; G. Brattman; D. Clifford W. Fitzpatrick A. Wickheim; J. E. V. and M. Goodwill; M. G. Shepard; J. E. Williams P. E. Wilford D. Wilson K. Kennedy, L. Dyke Unconfirmed Field notes Unconfirmed Photo; BCPM 371 Verbal description Verbal description Specimen; BCPM 14518 Photo; BCPM 372 Photo; BCPM 412 Field notes Field notes Specimen; BCPM 14519 Photo; BCPM 425 Verbal description Verbal description Field notes 88 THE CANADIAN FIELD-NATURALIST River, Denman Island, Lake Cowichan) where they often caught insects attracted to the animals and their droppings, and also probed manure piles for food items. Birds at Pachena Point and Nanaimo even foraged on lawns and in gardens. Both specimen records were from birds picked up dead and brought to the Provincial Museum for examination. The egrets weighed 212 gand 236 g, well below weights of 300-400 g cited by Palmer (1962). Also, the stomachs were empty, which suggests the birds may have died of starvation after long-distance flights. In summary, all records to date of the Cattle Egret in British Columbia are from the south coastal region, except for one sighting in the southern interior at Salmon Arm. The three 1970 records and three 1973 records could, in each case, conceivably have resulted from a single wide-ranging individual. At least five, but more likely six, birds were involved in the 1974 records, and at least four, but more likely five, in the 1975 records. All British Columbia records have been of single birds in late fall and early winter (extreme dates 15 November and 7 January). This closely follows the pattern of sightings in Oregon and Washington. The “Seasonal Reports” in Audubon Field Notes and American Birds list six records of Cattle Egrets for Oregon from 1965 to 1975, ranging in date from 22 November to 17 January, with a single record on | March 1975, which may involve a bird that success- fully wintered. In Washington, there have been five records (including an unconfirmed report in 1969), spanning the period 10 October to 8 January. In the absence of banding recoveries, the origin of British Columbia birds is uncertain, but the most likely route of dispersal is directly up the Pacific coast Vol. 91 from breeding areas in California or even in western Mexico. This hypothesis is supported by the fact that 6 of the 1! Oregon and Washington records are from the outer coast. But, even if the British Columbia birds came from the nearest Cattle Egret breeding colony at the Salton Sea in southern California, they must have travelled a minimum of 1850 km from where they were hatched. We thank the numerous observers who provided us with field notes, photographs, and specimens of Cattle Egrets; also Jerome A. Jackson, Anthony J. Erskine and J. E. V. Goodwill for their comments on the manuscript. Literature Cited Blake, C. H. 1961. Notes on the history of the Cattle Egret in the New World. Chat 25: 24-27. Campbell, R. W. and D. Stirling. 1971. A photoduplicate file for British Columbia vertebrate records. Syesis 4: 217-222. Campbell, R. W., M. G. Shepard, and R. H. Drent. 1972. Status of birds in the Vancouver area in 1970. Syesis 5: 137-167. Crosby, G. T. 1972. Spread of the Cattle Egret in the Western Hemisphere. Bird-Banding 43: 205-211. Davis, D. E. 1960. The spread of the Cattle Egret in the United States. Auk 77: 421-242. Palmer, R.S. 1962. Handbook of North American birds. Volume |. Yale University Press, London. 567 pp. Rice, D. W. 1956. Dynamics of range expansion of Cattle Egrets in Florida. Auk 73: 259-266. Small, A. 1974. The birds of California. Winchester Press, New York. 310 pp. Witzeman, J., J.P. Hubbard, and K. Kaufman. 1975. Southwest region. American Birds 29: 1014-1018. Received 14 June 1976 Accepted 14 October 1976 A Great Blue Heron Preying on Shiner Perch in Deep Water JEAN-GUY J. GODIN Department of Zoology, University of British Columbia, Vancouver, British Columbia V6T 1W5 Present address: Department of the Environment, Fisheries and Marine Service, Pacific Biological Station, Nanaimo, British Columbia V9R 5K6 Herons feed primarily on fish, and feeding methods common to all species are typically the “stand and wait” and “wade or walk slowly” behavior patterns (Lowe 1954; Meyerriecks 1960, 1962). Such aspects of feeding behavior are generally restricted to herons foraging in relatively shallow waters. I observed a Great Blue Heron (Ardea herodias L.) preying on shiner perch (Cymatogaster aggregata Gibbons) in deep water, and in this note I describe its feeding behavior and present quantitative estimates of its feeding efficiency. Observations and Discussion On the afternoon of 20 September 1975, a yearling Great Blue Heron (based on Palmer’s (1962) criteria) was seen landing ona floating platform near the wharf 1977 of the Pacific Biological Station, Departure Bay, British Columbia. Throughout the afternoon, the sky remained clear and sunny, the air temperature was about 20°C, and the water surface was calm. The plat- form (about 9 X 5 m), floating in about 10 m of water, was open except for a few planks extending across its length and width and suspended about 20 cm above the water surface. Before the heron left the wharf area, a total of 95 min of observation was made with the aid of stopwatches and a camera from within a docked boat about 7 m away from the platform. My presence did not appear to influence the heron’s activities. Approximately 5 min after its arrival, the heron began preying on fish swimming in the water enclosed by the platform. Since I could not see the fish from my vantage point, the heron’s feeding postures, described in the following paragraph, are assumed to have been displayed in response to the movements of the fish. The fish were later identified as yearling shiner perch (approximately 5-9 cm total length). Assuming a mean length of 7cm for the observed fish, an estimated mean wet weight of 5 g per individual was obtained, from Gordon’s (1965) length—weight relationships. The perch were aggregated (60-80 individuals) near (about 0-1 m) the water surface, feeding on plankton in shadows created by the overhead planks. When not walking on the planks in the “wade or walk slowly” or “low stalk” (Recher and Recher 1972) postures, the heron was generally observed to be in the “stand and wait” posture. The bird remained in this posture until prey was sighted. With its eyes fixed on the perch, the heron followed its movements with compensating eye and head movements but rarely moved other body parts. As the fish apparently approached to within | m or so of the bird, the latter’s long body axis slowly and gradually shifted down- ward and forward in the direction of the water surface with simultaneous uncoiling of the neck and bending of the legs. These gradual changes in head and body orientation are similar to those typically exhibited by feeding herons, excepting that the body of the bird was more steeply angled downward relative to the horizontal, owing to the higher position of the feet (20 cm above, rather than 0-20 cm below, the water surface). After the downward inclinations of its head and body, the heron either adopted a more erect posture or quickly exhibited a “strike” with its bill at the nearby prey. A very important factor in an animal’s feeding ecology, contributing to its survival and reproductive success, is feeding or foraging efficiency. Two com- monly used estimates of this efficiency are percentage capture success and food ration consumed per individual. During its 95-min feeding period, the Great Blue Heron had a 90.3% capture success (1.e., NOTES 89 captured and consumed 28 perch out of 31 “strike” attempts), and consumed 1.47 g of prey per minute. Krebs (1974) observed a capture success of about 41.0 + 3.5% (+ 1 SE) and a food consumption rate of about 1.5+0.15 g/min for solitary (non-flocked) adult Great Blue Herons feeding on a variety of intertidal fish species, including shiner perch. For aggregated (flocked) herons, he reported individual capture successes ranging from about 45.0 + 3.5 to 68.5+5.5% and food consumption rates ranging from about 1.55+0.15 to 5.15 +1.35 g/min. Recher and Recher (1969) reported a capture success of 64-75% with 0.3-0.7 g of prey consumed per minute for the Little Blue Heron (Florida caerulea). Later, Recher and Recher (1972) reported similar estimates (64-67% and 0.7 g of prey consumed per minute) for the same species, and a 48% capture success with 0.3-0.4 g of prey consumed per minute for the Reef Heron (Egretta sacra). A capture success of 90-97% (Siegfried 1971) associated with 0.13-0.51 g of prey consumed per minute (Siegfried 1972) has been documented for the Cattle Egret (Ardeola ibis). These values of feeding ef- ficiency were obtained for herons feeding in shallow waters, in pastures, or on flats.’ Although herons generally feed in shallow waters, some species have evolved specialized behavior patterns to prey on organisms in deep waters, especially when prey availability is high there (Meyer- riecks 1962). Diving or swimming (or both) after prey has been observed in the Common Heron, Ardea cinerea (Lowe 1954; Stacey and Gervis 1967), in the Green Heron, Butorides virescens (Meyerriecks 1960), and in the Great Blue Heron (Bent 1926). These deep-water feeding methods, however, do not appear to be very successful (Stacey and Gervis 1967). Palmer (1962) mentioned that the Great Blue Heron “occa- sionally fishes on the wing,” but failed to provide further details. In reviewing the literature on the Green Heron, Meyerriecks (1960) reported that this species may feed in deep water by remaining station- ary on structures such as logs, posts, or a rowboat, on which it either remains crouched with head and neck retracted or extends its head and neck downward toward the water, or alternatively by diving into the water from a plank, the edge of a park pool, or the shore of a pond or stream. He did not mention how successful these various feeding adaptations were. This paper is the first to my knowledge which presents quantitative estimates of the feeding effi- ciency of a Great Blue Heron feeding on fish in deep water. The behavioral postures reported here do not differ radically from those displayed by herons using the typical “stand and wait” feeding method while foraging in shallow waters, except for the greater downward head and body inclinations; these may be considered adaptive for feeding from structures 90 THE CANADIAN FIELD-NATURALIST elevated above the water surface. The food consump- tion rates of solitary Great Blue Herons reported by Krebs (1974) are very similar to the one presented here, but the 90.3% capture success of the heron feeding on perch in deep water is much greater than the capture successes of both solitary and aggregated herons feeding ona diversity of fish species in shallow, intertidal waters (Krebs 1974). Including the 90-97% capture success reported for Cattle Egrets foraging mainly on terrestrial arthropods (Siegfried 1971) and the high fish consumption rates reported for aggre- gated Great Blue Herons (Krebs 1974), the feeding efficiency values presented here appear to be the highest reported for Great Blue Heron and other heron species. These observations suggest that the Great Blue Heron can be an opportunistic and efficient predator. I thank P. A. Dill, A. J. Erskine, and W. S. Hoar for critically reading the manuscript. Literature Cited Bent, A. C. 1926. Life histories of North American marsh birds. United States National Museum Bulletin Number 135. 490 pp. Gordon, C. D. 1965. Aspects of the life-history of Cymato- gaster aggregata Gibbons). M.Sc. thesis, University of Vol. 91 British Columbia, Vancouver, British Columbia. 90 pp. Krebs, J. R. 1974. Colonial nesting and social feeding as strategies for exploiting food resources in the Great Blue Heron (Ardea herodias). Behaviour 51: 99-131. Lowe, F. A. 1954. The heron. Collins, St. Jame’s Place, London. 177 pp. Meyerriecks, A. J. 1960. Comparative breeding behavior of four species of North American herons. Publication of the Nuttall Ornithological Club, Number 2. 158 pp. Meyerriecks, A. J. 1962. Diversity typifies heron feeding. Natural History 71(6): 48-59. Palmer, R.S. (Editor). 1962. Handbook of North Ameri- can birds. Volume |. Yale University Press, New Haven, Connecticut. Recher, H. F. and J. A. Recher. 1969. Comparative forag- ing efficiency of adult and immature Little Blue Herons (Florida caerulea). Animal Behaviour 17: 320-322. Recher, H. F. and J. A. Recher. 1972. The foraging be- haviour of the Reef Heron. Emu 72: 85-90. Siegfried, W. R. 1971. Feeding activity of the Cattle Egret. Ardea 59: 38-46. Siegfried, W.R. 1972. Aspects of the feeding ecology of Cattle Egrets (Ardeola ibis) in South Africa. Journal of Animal Ecology 41: 71-78. Stacey, J. V. and G.R. Gervis. 1967. Heron apparently feeding in deep water. British Birds 60: 49-50. Received 6 August 1976 Accepted 27 September 1976 Spotted Redshank Sighted in Southern Ontario H. H. AXTELL,! P. BENHAM,? and J. E. BLACK3 'R.R. #1, Fort Erie, Ontario L2A 5SM4 2112 Linden Street, Kenmore, New York 14214 3Physics Department, Brock University, St. Catharines, Ontario L2S 3Al On 25 July 1976 we found and identified a Spotted Redshank (T7ringa erythropus) on a pond near Queenston, Ontario (the pond was closer to the small town of St. Davids, 43°10’N, 79°06’ W). W.E. Godfrey (1966. The birds of Canada. National Museum of Canada Bulletin 203. 428 pp.) cites a single hypothetical Canadian record for the Redshank (Tringa totanus) and notes that the bird might have been a Spotted Redshank. In a private communi- cation to the authors, Godfrey writes “There was no valid record of the Spotted Redshank for Canada until 1970. In the autumn of that year positively identifiable photos were taken of one at the Reifel Waterfowl Refuge, near Vancouver, B.C. (Campbell, R. W., M. G. Shepard, and R. H. Drent. Syesis 5: 151-152, 1971). There is also a sight record of one at the same locality in early May 1971 (American Birds 25(4): 791, 1971). Another sight record is from Newman Sound, Terra Nova National Park, New- foundland, where one was observed from May 15 to 21, 1974 (Finch, D. W. American Birds 29(1): 127, 1975). This specimen was in breeding plumage and therefore presumably unmistakable.” In view of the exceptional rarity of the Spotted Redshank in Canada, and the excellent opportunity we had to observe the bird, we present a summary of our visual observations in this note. For a more detailed account the reader is referred to a report submitted by one of us (H. H. A.) to the Buffalo Ornithological Society* and to the Ontario Orni- thological Records Committee.** *Buffalo Ornithological Society, Buffalo Museum of Science, Buffalo, New York, USA. **Sent to: C. E. Goodwin, 11 Westbank Crescent, Weston, Ontario. We understand that at least one other report on the Spotted Redshank was filed with this committee. 1977 The bird was first seen under clear skies at about 1100 hours in a telescope at 20-power. It was at a distance of about 140 m, and was in a settling pond operated by Canadian Canners. This pond is a new one and in places there is water to a depth of 8 or 10 cm, whereas in other places there is no water at all, only bare earth. A 3-m dike surrounds the pond which is about 275 m long and 120 m wide. About 100 Lesser Yellowlegs (Tringa flavipes) and 25 dowitchers (Lim- nodromus sp.) were present along with several hundred smaller shorebirds. The feature of the bird which first attracted our attention was a long bill with a red base. The bill length was about twice the size of the head and noticeably longer, in proportion to the head, than the bills of Lesser Yellowlegs in the same field of view of the telescope. The basal red extended a little less than half the bill length, and at 60-power a dusky strip on the top of the upper mandible was observed to parallel the red region. The bird was only slightly larger than the Lesser Yellowlegs, and when not standing beside a Lesser Yellowlegs was not separable from these by size alone. The bill was as slender, in proportion to the head, as the bills of the Lesser Yellowlegs. We were startled when the bird moved to shallow water and its red legs became visible. The color was easily seen at 20-power. In fact later in the day, when the bird was about 275 m from us, the leg color and the basal bill color could be seen with 40-power. A final striking field mark, visible in binoculars at 140 m, was the white which extended from the rump up the back ina manner similar to the pattern found in dowitchers. The white was seen on several occasions when the bird flew, and on at least one occasion when the bird preened. It was clearly visible when the bird left the pond ‘at about 1700 hours. (Although a number of people have looked for the bird since that time, it has not been reported at St. Davids.) Before Fieldfare in Ontario! DAVID J. T. HUSSELL and MICHAEL J. PORTER NOTES 91 leaving the pond, the bird flew over it at a height of about 9m repeatedly uttering a call that some observers said resembled a Semipalmated Plover (Charadrius semipalmatus) call. If one accepts the clearly visible field mark of the red legs then the only birds known in North America that could qualify as the bird seen are the Redshank and Spotted Redshank. There are several field marks that support the latter choice. First, there was no sign of the white hind edges of the wings said to characterize the Redshank. Second, underparts of the bird were very much darker than those depicted for the Redshank in A field guide to the birds of Britain and Europe (Peterson, R.T., G. Mountford, and P. A. D. Hollom. 1966. Houghton Mifflin Company, Boston. 344 pp.). There was considerable dark mottling on the underparts of the bird. Finally, there was the long bill, substantially longer in proportion to the head than the bills of the Lesser Yellowlegs; this one expects for the Spotted Redshank. Although a very heavily oiled bird might not show the white hind edges of the wing, and would show overall dark appearance, it might also have oil on the rump and back that was not observed. The plumage, in fact, seemed to be one in transition between the summer and winter phases of Spotted Redshanks depicted in the field guide. Although we believe in general that a good photo- graphic record should be given more credence than visual observations, the field marks of this particular bird, and the excellent opportunity we had for careful and lengthy observation, make this record one of the most credible of visual observations. At least seven other adult bird watchers saw the bird, and all easily saw the red legs. Received 9 August 1976 Accepted 24 October 1976 Long Point Bird Observatory, P.O. Box 160, Port Rowan, Ontario NOE 1M0 On the morning of 24 May 1975, M. J. Porter heard the distinctive calls of a Fieldfare (Turdus pilaris) at the Long Point Bird Observatory’s station on Courtright Ridge, Long Point, Ontario (42°33’ N, 'A publication of the Long Point Bird Observatory. 80°17’ W). He saw the bird in the treetops and, being familiar with the species in Europe, was able to identify it immediately. Half an hour later Alex Steele found the Fieldfare caught in a mist net which he and Porter had set alongside some spruce and pine trees. Plumage details were recorded and it was banded, 92 THE CANADIAN FIELD-NATURALIST weighed, and measured. The bird was then taken to the mainland, where it was photographed (Figure 1) and examined in greater detail by D. J. T. Hussell and several other people, including M. Bradstreet, E. H. Dunn, G. W. Miller, J. Woodford, and P. S. Wood- ford. When it was released later the same day, the Fieldfare called and flew to a nearby tree, where it was watched for a few minutes. It was not seen again. The bird was a large thrush, similar in size and build to an American Robin (Turdus migratorius), and identified by its gray head and rump, chestnut back, and speckled breast. The wing chord measured 139 mm, indicating that it was probably a female (males 140-153 mm, females 135-148 mm, Witherby et al. 1940). A detailed description, including measurements, and copies of color slides have been deposited with the Ontario Ornithological Records Committee at the Royal Ontario Museum. There was no indication that the bird had recently FIGURE |. Fieldfare, Long Point, 24 May 1975. Fromacolor slide by Isabel Smaller. Vol. 91 been in captivity. The legs, feet, and wings were examined carefully for wear and found to be in good condition. The tips of the tail feathers were somewhat worn, but the ends of the shafts were intact, project- ing slightly beyond the webs. The summer range of the Fieldfare extends from south-central Siberia west to Scandinavia and south to Switzerland and Hungary, with recently estab- lished outpost populations in southwestern Green- land (Voous 1960) and the British Isles (Williamson 1975). Fieldfares winter mainly in southern and central Europe, including the British Isles (Witherby et al. 1940). Previous North American records include speci- mens from Stamford, Connecticut (April 1878) and Jens Munk Island in Foxe Basin, NWT (1939); a photograph from St. John’s, Newfoundland (1 January 1973); sight records at Ottawa, Ontario (8 January 1967), Bombay Hook, Delaware (30 March-4 April 1969), and Larchmont, New York (3-11 February 1973) (Finch 1973). The origin of these North American birds is a matter of speculation. Salomonsen (1951) says that the Greenland breeders are sedentary, implying that they do not migrate east across the Atlantic to winter in the Old World. The recent records suggest the possibility of the develop- ment of a pattern of regular wintering in North America by a portion of the Greenland population. The present record is the first for Ontario to be documented with material evidence (photograph), and as such it constitutes an addition to the Ontario list accepted by the Ontario Ornithological Records Committee (C. E. Goodwin, personal communica- tion). We thank the members and employees of the Long Point Company for facilitating the operation of the Observatory’s station on Courtright Ridge. A. D. Brewer and C. E. Goodwin commented on an earlier draft of the manuscript, and D. W. Finch provided information on previous North American records. Literature Cited Finch, D. W. 1973. Northeastern Maritime Region. Amer- ican Birds 27: 586-591. Salomonsen, F. 195]. The immigration and breeding of the Fieldfare (Turdus pilaris L.) in Greenland. Proceedings of the 10th International Ornithological Congress. pp. 515-526. Voous, K. H. 1960. Atlas of European birds. Nelson, New York. 284 pp. Williamson, K. 1975. Birds and climatic change. Bird Study 22: 143-164. Witherby, H. F., F. C. R. Jourdain, N. F. Ticehurst, and B. W. Tucker. 1940. The handbook of British birds. Volume II. Witherby, London. 368 pp. Received 28 May 1976 Accepted 20 August 1976 1977 NOTES 93 Genetic Variants in Canada of the Rainbow Trout, Salmo gairdneri, Called Golden Trout and Palomino Trout ROBIN E. CRAIG! and E. J. CROSSMAN2 1Ontario Ministry of Natural Resources, Midhurst, Ontario LOL 1X0 2Royal Ontario Museum, Toronto, Ontario M5S 2C6 Prior to 1974 specimens of an interesting variety of rainbow trout (Salmo gairdneri), generally referred to by anglers as “golden trout,” had been irregularly taken in USA waters of Lake Erie and the Buffalo- Niagara River area of New York State(R. B. Kenyon, personal communication). These unique trout are not to be confused with the true golden trout, Salmo aquabonita Jordan. “Golden trout” were first re- corded from Ontario waters in 1974. The first specimens reported were taken by anglers from the Upper Niagara River just upstream from the city of Niagara Falls. These fish were marked with a left pelvic-fin clip. The origin of these variants was traced to Penn- sylvania. They had been reared and released into Lake Erie and its tributaries, as part of a cooperative program involving the Pennsylvania Fish Commis- sion and the 3CU Trout Association (a sportsmen’s group). Two rainbow variants had been reared and released. These were known as “golden trout” (golden-orange in color, without black spots) and “palomino trout” (golden in color with a reduced number of faint black spots as compared with normal rainbow trout). In 1975 at least eight variant trouts were taken in Ontario waters: May, two golden trout incommercial nets from Lake Ontario, 1.6 km (1 mi) west of the mouth of Niagara River, left pelvic clips; May, one palomino trout in commercial nets from Lake Ontario, near Grimsby, Pennsylvania, tag; September, one golden trout in commercial nets from Lake Erie, 1.6 km (1 mi) east of Port Dover, left pelvic clip; October, one golden trout and one palomino trout, Ontario Ministry of Natural Resources crew, Ontario Hydro diversion canal, Niagara Falls, left pelvic clip. The tagged individual caught in Lake Ontario near Grimsby had been hatched in May 1972. It had been released (at length of 175-225 mm) into Lake Erie off the mouth of a tributary stream, Trout Run, on 4 April 1975. The Pennsylvania records referred to this individual as a “Palomino Steelhead Rainbow Trout.” The name golden trout as used by the anglers was derived from the unique color, since the anglers were unable to recognize them as rainbow trout, Sa/mo' gairdneri. “Golden trout” was also the name given these fish in the original genetic experiment from which they were developed. The golden color is a combination of genetic effects on background color, resulting from color dilution because of an absence of melanophores. The “golden” trout are basically orange-gold (O-17-10°) (see Villalobos-Dominquez and Villalobos 1947) on the back. The sides are golden to silvery with a darker orange lateral band (SO-17-9°). The opercles and cheeks are reddish orange (SSO-14-10°), and the lower sides and belly milk white. The dorsal, adipose, and caudal fins are yellow. The anal fin appears to lack color. The paired fins are yellow with an orange leading edge. The “palomino” trout have the same overall coloring as the “golden” trout (sometimes darker) but show a pattern of black spots on the head, back, and caudal fin. The caudal spotting is more predominant on the upper lobe. The spotting is much reduced (number, size, and density) from that of normal rainbow trout. The palomino trout seen in Ontario appear to have more spots than indicated by the illustration of that variant given by Wright (1972). Values for almost all morphometric characters for those examined fell within the central range of the values for Salmo gairdneri. The exception was gill rakers which were at the lower limit (9 on upper, 10-11 on lower). The specimen taken near Port Dover in September 1975 was a male with greatly enlarged white testes. Maximum total length of those seen has been 45.3 cm and maximum weight 1.15 kg at age I+. At least some individuals appear sexually mature at this size and age. According to Wright (1972) this “golden” trout was developed and popularized in 1963 as “West Virginia’s Centennial Golden Trout.” The variant arose from a single female, spawned in the fall of 1954, with a body pattern which was a mosaic of those of the “golden” trout and normal rainbow trout (see Clark 1970). This mosaic pattern has been referred to as “chimera.” When a chimera female is crossed with a normal male rainbow trout, the light- colored offspring with reduced spotting called “pal- omino” result. They are heterozygous for a gene causing color dilution. When two “palominos” are mated the progeny consist of normal pattern, pal- omino pattern, and golden pattern, in a precise ratio (1 normal: 2 palomino: | golden). 94 THE CANADIAN FIELD-NATURALIST The presence of these variants in Lake Ontario indicates an extension of range from Lake Erie. The route the fish took is unknown. Their earlier presence in the upper Niagara River would suggest passage over Niagara Falls. The capture at Grimsby might suggest the alternative route through the Welland Canal. The captures in the Ontario Hydro canal suggest that as a possibility. They would have to pass through the power generating equipment but Hydro officials say adult fish can pass and live. Presently it would appear the source of the variants is the Pennsylvania introductions. There has been, however, notice of 100 000 golden trout produced in Wisconsin by an organization called Salmon Un- limited (Anonymous 1975). If some of these were released in the Great Lakes or tributaries they could contribute to those which might be seen elsewhere in Canada. The possible ecological effects of the variants on rainbow trout in the lower Great Lakes are unknown. Major determinants of this effect and of the per- manence of the stock of variants are the rate of con- tinued introduction and degree of crossing after release. Since the variants react similarly to the wild rainbow phenotype (R. B. Kenyon, personal com- munication), spawning of variants with wild normal trout can be expected. According to Wright (1972), if a golden variant crosses with a normal rainbow, palomino trout result. If palomino trout mate with normal rainbows one half of the offspring are palominos and one half normal rainbows. Even if Vol. 91 palominos mate, one quarter of the offspring are normal trout (+ 1/2 palomino and 1/4 golden). Only when palomino trout cross with golden trout (= 1/2 golden and 1/2 palomino) or golden trout cross with golden trout (= all golden) is there no dilution of the variants. There are, therefore, at least two new forms of trout to add to the problem of identification of salmonids in the lower Great Lakes. The choice of terminology is unfortunate since golden trout is the accepted name for Salmo aqua- bonita, a species native to the western USA and introductions of that species have been attempted elsewhere in Canada. The authors acknowledge the cooperation and assistance provided by E. Ball, R. Borecky, R. H. Brown, B. Campbell, J. Cossitt, A. Giesche, R. B. Kenyon, P. J. MacDonald, J. W. Meade, III, and S. J. Nepszy. The color and morphometric descrip- tions were prepared by Cheryl Goodchild. Literature Cited Anonymous. 1975. Ontario fisherman and hunter. Out of Doors, October. p. 5. Clark, F. H. 1970. Pleitropic effects of the gene for golden color in rainbow trout. Journal of Heredity 611: 8-10. Villalobos-Dominquez, C. and J. Villalobos. 1947. Colour atlas. El] Ateneo, Buenos Aires. 46 pp. Wright, J. F., Jr. 1972. The palomino rainbow trout. Penn- sylvania Angler 41(3): 8-9, 26. Received 5 March 1976 Accepted 27 July 1976 Trauma-induced Paralysis in a Moose Calf GORDON A. CHALMERS! and MORLEY W. BARRETT2 1Alberta Department of Agriculture, Veterinary Services Division, Lethbridge, Alberta T1J 3Y5 2Alberta Department of Recreation, Parks and Wildlife, Fish and Wildlife Division, Lethbridge, Alberta T1H OH5 The progressive encroachment of man into wildlife habitats is a profound threat, individually and collectively, for many wildlife species today. We report, as an example, the fracture of a cervical vertebra and consequent paralysis in a free-ranging moose (Alces alces). A live 5- to 6-month-old female moose calf was found lying on its right side adjacent to a three-strand barbed wire fence in the Porcupine Hills of south- western Alberta; the animal was transported to the laboratory for further clinical examination and necropsy. Clinically the calf was markedly hyper- sensitive over the anterior thoracic and cervical vertebrae with reduced sensation at the coronary bands of all four limbs. The lids of the right eye were bruised and there was mild corneal opacity. Rectal temperature was 38.3°C. Blood for hematological examination was drawn by jugular venipuncture into Vacutainer** tubes containing potassium ethylene- diaminetetra-acetate. The calf was subsequently killed by an intravenous injection of Lethal. + At necropsy, the carcass was emaciated, with serous atrophy of fat depots. No gross lesions were observed in thoracic or abdominal organs or in the large arteries of the thorax and neck. The rumen contained a small volume of firm dry forage. A few flakes of fibrin were **Becton, Dickinson and Company, Canada Ltd., Missis- sauga, Ontario. *Haver-Lockhart, Calgary, Alberta. 1977 NOTES 95 FIGURE 1. Sagittal section through the axis and C,, demonstrating the fracture, hemorrhage, and upward displacement of fragments. The atlas has been removed. present in the right coxofemoral joint; synovial fluid was clear. No gross lesions were detected in the brain. Examination of the vertebral column, however, revealed that the third cervical vertebra was fractured, displaced dorsally, and impinged on the spinal cord (Figure 1). Yellow to brown discoloration of the cord and meninges was marked in this area. Histological examination of the affected spinal cord revealed dorso-ventral compression, severe distortion, and hemorrhage with widespread axonal swelling and extensive softening. Vascular endothelial swelling and proliferation were prominent findings. Hematological examinations provided the follow- ing data: hemoglobin, 14 g/100 ml; leukocytes, 4000/mm3; segmented neutrophils, 46%; lymph- ocytes, 50%; monocytes, 4%; sedimentation rate, 57 mm/h and 60 mm/24 h. Discussion The loss of forested land because of clearing for agricultural purposes has been identified as a major loss of moose habitat in Alberta (Lynch, G. M. 1974. Moose management plan for Alberta. Alberta De- partment of Recreation, Parks and Wildlife, Fish and Wildlife Division. Mimeo. 32 pp.). The increasing network of fences in forested areas may be an additional hazard to big-game animals, by altering or occasionally preventing, normal movement patterns of moose and, perhaps elk (Cervus canadensis). Moose and elk calves in west central Alberta have been observed caught in fences or separated from the maternal parent; attacks by farm dogs are often complicating factors (G. M. Lynch, personal com- munication; W. M. Samuel, personal communica- tion). R.L. Peterson (1955. North American moose. University of Toronto Press, Toronto. 280 pp.) reported that moose seldom attempt to jump, but when forced to do so, clear obstacles by rearing, placing the front legs over, and then springing or diving over these obstacles with their hind legs. Fences tend to be formidable barriers to calves when their ability to clear obstacles is less well developed than that of mature animals. Circumstantial evidence in this case suggested that the calf had attempted unsuccessfully to jump the fence. The assumption was supported by the proximity of the animal to the fence and by the nature of the injury. By failing to clear the fence effectively, the calf may have fallen heavily on its head, neck, or brisket. The ocular and palpebral lesions, serous atrophy of fat depots, and the presence of dry firm rumen contents indicated that the animal had been recumbent for several days prior to discovery. The authors thank B. Arnold, Alberta Fish and Wildlife Division, Claresholm, for locating and trans- porting the moose to the laboratory. J. Wood, V. Wong, and P. Mills of the Veterinary Services Division, Lethbridge, provided much appreciated laboratory technical assistance. Received 14 April 1976 Accepted 19 July 1976 96 THE CANADIAN FIELD-NATURALIST Vol. 91 A Recent Record of the Meadow Jumping Mouse, Zapus hudsonius, in the Northwest Territories RICHARD J. DOUGLASS! and ANDREW E. L. McNAUGHTON2 ‘Renewable Resources Consulting Services Ltd., 11440 Kingsway Avenue, Edmonton, Alberta T5G 0X4 Museum of Zoology, Department of Biology, Carleton University, Ottawa, Ontario KIS 5B6 A meadow jumping mouse was captured on 12 August 1975 during field studies in the Northwest Territories. The single specimen (skin and skull) was identified as Zapus hudsonius by C. G. van Zyll de Jong, and deposited in the National Museum of Natural Sciences, Ottawa, as NMC 42852. The animal was collected on the edge of a small thermokarst lake near the northwest end of Chick Lake (65°53’ N, 128°12’ W; Figure 1), 90 km north- west of Norman Wells, Northwest Territories. Hall and Kelson (1959) and Banfield (1974) considered the northern limit for the jumping mouse in the North- west Territories to be south of Great Slave Lake and the Mackenzie River. Krapu and Traugher (1972) recorded a specimen near the north shore of Great Slave Lake, 40 km northwest of Yellowknife. (The latitude was incorrectly published as 66°33’ N. The correct latitude for their specimen is 62°33’ N; Krapu, 1976, personal communication.) The Chick Lake capture extends the known range approximately 360 km. The vegetation at the present capture site consisted primarily of sedges (Carex spp.) and grasses (Calama- grostis spp.), with clumps of willow (Salix spp.). The forest surrounding the lake was dominated by black spruce (Picea mariana) with some white spruce (P. glauca) and larch (Larix laricina) intermixed. A relatively high degree of thermokarst erosion adjacent to this small lake was the only apparent feature that distinguished it from others in the area. The one meadow jumping mouse taken represented less than 0.005% of the total captures in 95 800 trap- nights expended at Chick Lake during the summers of 1973, 1974, and 1975. The most common species trapped were the northern red-backed vole (Cleth- rionomys rutilus) and the meadow vole (Microtus pennsylvanicus). Other species trapped at Chick Lake were, in descending order of frequency, chestnut- cheeked vole (Microtus xanthognathus), ermine (Mustela erminea), red squirrel (Tamiasciurus hud- sonicus), masked shrew (Sorex cinereus), heather vole (Phenacomys intermedius) (Douglass and McDonald 1976), arctic shrew (S. arcticus; NMC 42840), and pigmy shrew (Microsorex hoyi; NMC 42851). Other mammals sighted were varying hare (Lepus ameri- canus), muskrat (Ondatra zibethicus), marten (Martes americana), mink (Mustela vison), wolf (Canis lupus), black bear (Ursus americanus), moose (Alces alces), and caribou (Rangifer tarandus). These collections and observations were made during field studies conducted for Canadian Arctic Gas Study Limited by Renewable Resources Consult- ing Services Limited. Literature Cited Banfield, A. W. F. 1974. The mammals of Canada. Univer- FiGURE 1. Canadian and Alaskan distribution of the meadow jumping mouse, Zapus hudsonius, after Hall and Kelson (1959). @ Chick Lake capture, o Krapu and Traugher (1972), @ Youngman (1975). iy) sity of Toronto Press, Toronto. 438 pp. Douglass, R.J. and D. McDonald. 1976. A northern record for the heather vole, Phenacomys intermedius, in the Northwest Territories. Canadian Field-Naturalist 90: 82-83 Hall, E. R.and K. R. Kelson. 1959. The mammals of North America. Volume 2. Ronald Press, New York. 623 pp. Krapu, G. L. and D. L. Traugher. 1972. A recent record of NOTES 97 the meadow jumping mouse, Zapus hudsonius, in sub- arctic Canada. American Midland Naturalist 88(2): 467. Youngman, P. M. 1975. Mammals of the Yukon Territory. National Museums of Canada, Publications in Zoology, Number 10. 192 pp. Received 16 June 1976 Accepted 9 August 1976 Swarming of Dragonflies Noted at Drag Lake, Ontario T. E. Perry, M.S. PERRY, and J. E. K. PERRY Chagrin River Road, Gates Mills, Ohio 44040 We spent three vacation days at Drag Lake, near Haliburton and Minden, Ontario, 23, 24 and 25 June 1975. A few more than the usual numbers of dragon- flies were noted as we approached the lake from the south, but not as thick a swarm as at the lake itself. A specimen of Cordulia shurtleffi Scudder was flying about in downtown Minden, crowds and traffic not- withstanding. Roads to Drag Lake were filled with Libellula julia Uhler, squatting on bare ground, in both sun and shade. Drag Lake is situated near Haliburton, Ontario, about 50 km south of the main portion of Algonquin Park, 45.5°N, 78.1° W. The lake is about 1.5 X 6.5 km at its widest and longest points and is said to be as deep as 200 m. During the late nineteenth century extensive lumbering operations consisted of dragging logs across the lake in winter, hence the name Drag Lake. The lake’s shores are boulder-strewn and irregular (roughly Y-shaped) in outline; the drop-off appears steep in most places. Inland are numerous small, shallow bays filled with aquatic vegetation, providing favorable dragonfly breeding habitats. Weather conditions at Drag Lake on 23 through 25 June were excellent for observing and collecting insects. Day-time temperatures were warm (about 23°C), sunny, with slight-to-moderate wind. Pre- cipitation consisted of a very small amount of rain during the trip up to the lake area, just enough to dampen exposed surfaces. The dragonfly swarm seemed concentrated at Drag Lake. We explored other areas near and approaching the lake, but found dragonflies much more generally dispersed. The swarm was stationary and was feeding; it consisted of about 10 species of Odonata. Vertical stratification of the swarm was observed (Table 1). Mid-level (1 through 4 metres) was the most active area for both numbers of species and individuals. The absence of certain common types of dragonfly TABLE 1—Dragonfly species present, their observed swarming height, and frequency at Drag Lake, Ontario on 23-25 June 1975 Low Species present (0-1 m) Cordulia shurtleffi Scudder Epitheca cynosura (Say) Epitheca princeps Hagen Epitheca spinigera Selys Leucorrhinia frigida Hagen Leucorrhinia hudsonica (Selys) Libellula julia Uhler Libellula quadrimaculata Linne Macromia illinoiensis Walsh Somatochlora williamsoni Walker mm Swarming height Mid High (1-4 m) (> 4m) Frequency x common x common Xs rare x most common occasional occasional common x common X x rare x rare 98 THE CANADIAN FIELD-NATURALIST behavior within the swarm was interesting to note. Competition in form of territorial aggressiveness and defence and also reproductive activities were not observed. The dragonflies seemed also not as wary of humans as often they are. The mosquito season was in full course; as we walked through the dragonfly swarm at evening the dragonflies came closer to us, flew at lower levels, and fed on the mosquitoes attracted by our presence. Also interesting to note was the scarcity of damselflies at Drag Lake: individuals seen numbered less than 10 orso. It is possible that the large number of dragonflies consumed many damsel- flies earlier. Mosquitoes, however, appeared the swarm’s major target. As to space and time factors, swarming continued throughout each of the three days, and was heaviest at midday to near sundown. Swarming occurred entirely over land rather than water surfaces. Some dragonflies were not part of the swarming phenomenon at Drag Lake but were taken as part of our sampling. Along the lake shore were Gomphus spicatus Hagen and Basiaeschna janata (Say). At one localized and protected cell on the lake’s northwest side we found Aeshna canadensis Walker clinging to tree branches and trunks at the forest edge, a habitat noted by Walker (1958). Didymops transversa (Say) was also taken on vegetation along roadways. Dragonfly swarming has been studied and noted in the literature. “Swarm-feeding” may be the best term describing our observations at Drag Lake. Corbet (1963) discussed this in some detail. Corbet also commented on the lack of interaction among dragon- flies feeding upon other insect swarms. Kormondy (1959) observed six different species of dragonfly flying together in Michigan, with Epitheca spinigera Vol. 91 Selys in the majority; this is similar to our obser- vations at Drag Lake. Walker and Corbet (1975) also mentioned spinigera’s appearance in numbers in southern Ontario. None of the dragonflies described here are con- sidered rare for the locality, and may be considered well within their given ranges (Needham and Westfall 1955; Walker 1958; Walker and Corbet 1975). Acknowledgments Thanks are given to Miriam Ouellette, proprietress of Drag Lake Resort, for permission to study and collect the dragonflies. Special thanks are also given George Bachmann of the Social Studies Department, Memorial School, Mentor, Ohio, for information on the geography of the Drag Lake region. Literature Cited Corbet, P.S. 1963. A biology of dragonflies. Quadrangle Books, Chicago, Illinois. 247 pp. Kormondy, E. J. 1959. The systematics of Tetragoneuria, based on ecological, life history, and morphological evidence (Odonata: Corduliidae). Miscellaneous Publica- tions, Museum of Zoology, University of Michigan 107: 1-79. Needham, J. G. and M. J. Westfall. 1955. A manual of the dragonflies of North America (Anisoptera). University of California Press, Berkeley. 615 pp. Walker, E. M. 1958. The Odonata of Canada and Alaska. Volume Two, Part III. University of Toronto Press, Toronto. 318 pp. Walker, E. M. and P.S. Corbet. 1975. The Odonata of Canada and Alaska. Volume Three, Part III. University of Toronto Press, Toronto. 307 pp. Received 28 May 1976 Accepted 11 August 1976 Two Recent Bobcat (Lynx rufus) Specimens from Southern Ontario DAVID NAGORSEN and RANDOLPH L. PETERSON Mammalogy Department, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6 As a result of a recent survey of bobcat (Lynx rufus!) material in major Canadian institutions, we found that two specimens of this species in the collec- ‘Recently, authors such as van Zyll de Jong (1975) con- sidered Lynx as a subgenus of Felis rather than a separate genus and assigned the lynx and the bobcat to the genus Felis. Yo avoid confusion with the nomenclature used by Peterson and Downing (1952), Peterson (1966), and Banfield (1974), we used the genus Lynx for the bobcat in this paper. tions of the Royal Ontario Museum (ROM) appar- ently represent the only museum specimens that have been collected in southern Ontario since 1906. One specimen (ROM 25497) is an adult male from Pakenham Township, Lanark County, 1953. The other (ROM 67947) obtained from the Fish and Wildlife Research Branch of the Ministry of Natural Resources, is a young male, found as a road kill on Highway #11, Muskoka Township, Bracebridge District, 1972. These two specimens are of considerable interest 1977 since the racial affinities of the present southern Ontario bobcat population are questionable. Peter- son and Downing (1952) assigned the original popula- tion to the subspecies L. r. gigas, a race that is found also in New Brunswick, Nova Scotia, and possibly southern Quebec (Peterson 1966). But since the only gigas material from Ontario available to Peterson and Downing (1952) was collected prior to 1906, these authors suggested that L. r. gigas may have become extinct in southern Ontario within the last half- century, and may have been replaced by some other race. According to the distribution map of Peterson and Downing (1952), it is possible that either of two races of L. rufus could have recently extended their range into southern Ontario: L. r. rufus, found in the eastern United States, or L. r. superiorensis from the northwestern Great Lakes region. Although Banfield (1974) reported that L. r. rufus has recently invaded southern Ontario, this conclusion apparently was not based on recent material. We have compared the skulls of the two recent ROM specimens with those of gigas, rufus, and superiorensis and when the diagnostic characters described by Peterson and Downing (1952) and Peterson (1966) are used, the skulls appear to conform closely to L. r. superiorensis. Both skulls differ from those of L. r. rufus in their dorsal contour and by their relatively smaller third upper premolar (PM)?). ROM 25497 could also be distinguished from skulls NOTES 99 of gigas by its relatively wider and shorter palate and the ratio of its maxillary tooth row to the width of the palate. Unfortunately, the palatal width of ROM 67947 could not be measured because the skull is partially damaged. The skull of this specimen, however, does conform closely to young male material of superiorensis. ROM specimen records and reports from trappers indicate that the populations of L. r. superiorensis in Ontario are the result of two recent invasions. Probably derived from the population of superioren- sis in Minnesota, this race of bobcat first appeared in Ontario early in the century in the region west of Lake Superior, where it rapidly expanded its range north and east. In the late 1940s, a second invasion occurred when L. r. superiorensis crossed into Ontario from the upper peninsula of Michigan, and by 1952 specimens had been obtained from Sault Ste. Marie (formerly Algoma) District and Cockburn Island, Espanola (formerly Manitoulin) District (Peterson and Down- ing 1952). Although inconclusive, the identification of the two recent specimens from Lanark County and Bracebridge District as superiorensis suggests that this western race has now spread into southern Ontario (Figure 1). Fur harvest records obtained from the Fish and Wildlife Research Branch, Ministry of Natural Resources for 1973-1975, show that eight bobcats were taken in southern Ontario for this period with specimens trapped in the following southern | EE ae SEE SY FicuRE |. Locations of two recent bobcat (Lynx rufus) specimens from southern Ontario and the distribution of the western race superiorensis. Stippling represents the approximate range of superiorensis in Ontario, each dot represents one or more specimens in the Royal Ontario Museum, and squares represent the two specimens from southern Ontario. 100 Ministry districts: Lindsay, Cambridge, Tweed, Lanark, Pembroke, and Minden. In order to confirm the taxonomic status of this population of L. rufus now inhabiting southern Ontario, the Royal Ontario Museum is attempting to acquire specimens from this region. In a recent study applying numerical tax- onomy, van Zyll de Jong (1975) analyzed skull and body ratios of the three subspecies of bobcat dis- cussed in this paper and found that they show intra- specific differences. It would be interesting to com- pare the present population of L. rufus in southern Ontario with these three subspecies using modern taxonomic methods if, and when, an adequate sample size can be obtained. We thank Bruce Stephenson, Fish and Wildlife Research Branch, Ministry of Natural Resources, for reviewing his bobcat records and providing us with THE CANADIAN FIELD-NATURALIST Vol. 91 the Ontario fur harvest records. Literature Cited Banfield, A. W. F. 1974. Mammals of Canada. University of Toronto Press, Toronto. 438 pp. Peterson, R. L. 1966. The mammals of eastern Canada. Oxford University Press, Toronto. 465 pp. Peterson, R. L. and S.C. Downing. 1952. Notes on the bobcats (Lynx rufus) of eastern North America with the description of a new race. Contributions of the Royal Ontario Museum of Zoology and Palaeontology, Number 33. 23 pp. van Zyll de Jong, C. G. 1975. Differentiation of the Canada lynx, Felis (Lynx) canadensis subsolana, in Newfound- land. Canadian Journal of Zoology 53: 699-705. Received 7 May 1976 Accepted 16 June 1976 News and Comment Note to Authors and Referees For future submissions to The Canadian Field- Naturalist, 1 encourage all authors to suggest poten- tial referees. Although these referees may not be selected to review a particular author’s manuscript, they willadd to my list from which referees are chosen. This request is now incorporated into the updated Instructions to Contributors. Editors are constantly searching for experts in particular fields of study to review manuscripts. These experts should base their acceptance or rejection of manuscripts on sound scientific reasoning and on the significance of the contribution to the ever-expanding literature. Authors are always free to rebut the criticisms of referees and editors but their refutations should be based on sound arguments. Referees are particularly requested to keep the average reader in mind when they are evaluating manuscripts. It is important that the research results, and interpreta- tions from them, are communicated ina logical, clear, and interesting manner to the greatest possible number of our readers. My current editorial policy is to let referees make their own personal choice whether to remain anon- ymous or whether, by signing their comments for the author, to reveal their identity. From time to time the question of the anonymity of referees is debated (see “Towards open refereeing” by Peter Robertson in the New Scientist 71(1014): 410, 1976) and both benefits and disadvantages are recognized. Certainly if a referee reveals his identity, his reputation as well as his relationship with the author is at stake. Editors are particularly grateful that many referees, as a duty to the scientific community, spend a considerable time and effort to perform a service that brings but little reward other than personal satisfaction for a job well done. All referees should try to show humane considera- tion for authors (see “A plea for tolerance and courtesy” by David E. Davis in BioScience 26(3): 171, 1976). If this is done, then even a negative evaluation, if accompanied by constructive comments and softened by a word of encouragement, can promote good relationships. A working arrangement beneficial to all is our aim. LORRAINE C. SMITH, Editor A New Series on the Biological Flora of Canada We feel that the time is right to initiate a series on the Biological Flora of Canada wherein ecological life history information for important members of our flora is assembled. Sucha series, analogous to, but not identical with, that of the Journal of Ecology was in The Canadian Field- Naturalist proposed by George H. La Roi and he will be elaborating on the subject in a forthcoming editorial. In order to start the series off on the right foot, it will be necessary to consult with others and to deliberate with care regarding the guidelines and final format. Environmental Concerns I draw the attention of our readers to two recent thought-provoking and interrelated editorials: “The impact statement boondoggle” by D. W. Schindler, published in Science 192(4239): 509, 1976 and “Where is Canadian environmental science headed?” by D. E. Sergeant, published in the Canadian Society of Environmental Biologists Newsletter/ Bulletin 33(3): 3-5, 1976. Dr. Schindler notes that “impact statements seldom receive the hard scrutiny that follows the publication of findings in a reputable journal.” Furthermore, he notes that “Having seen the results of many of these impact studies, and evaluated proposals for second- generation studies, I believe the idea has backfired.” Dr. Sergeant also points out some of the problems in the current government approach to Canadian science—decline in funds for research by government and universities, and an increased reliance on con- sulting companies. The latter do not publish their research through normal scientific channels and, therefore, many of their reports will eventually be lost. The views of these concerned scientists should be read by ecologically concerned citizens and hopefully heeded by government policy makers. LORRAINE C. SMITH, Editor 101 102 Herpetology Information Needed I am studying the status and ecology of the Blue Racer, Cricket Frog, Island Water Snake (of the Lake Erie Islands), Spiny Softshell Turtle, Queen Snake, and Fowler’s Toad in Ontario. The Canadian Am- phibian and Reptile Conservation Society, the Ontario Ministry of Natural Resources, and the THE CANADIAN FIELD-NATURALIST Vol. 91 Canadian Wildlife Service are also concerned with these animals and have some information about them, but any additional information on their occurrence, habitats, or changes in number would be most helpful. Please send information to Craig Campbell, 421 King Street North, Waterloo, Ontario N2J_ 3Z4. Northeastern Regional Meeting of the Animal Behavior Society Memorial University of Newfoundland will host the 1977 Northeastern Regional Meeting of the Animal Behavior Society in St. John’s, Newfound- land. The conference will be held during October at St. Bride’s College in St. John’s where room and board will be available at reasonable rates. Papers (15 to 20 min) and poster presentations from all areas of animal behavior are invited. Morning plenary sessions will focus on different groups of marine organisms (invertebrates, fish, birds, and mammals) and will address themselves to three topic areas—navigation, communication-social be- havior, and behavioral development. A film session Third Annual Ontario Ecological Colloquium This colloquium will be held on 18 and 19 April 1977 at The University of Western Ontario, London, Ontario. It is sponsored by the Departments of Plant Sciences and Zoology. The organizers hope that as many Ontario ecolo- gists as possible will present papers about their past and present research or initiate discussions on matters of common interest and concern. Tentative session titles are avian ecology; mammalian ecology; fresh- on marine mammals and a discussion on constraints on learning are also planned. There will be ample opportunity to tour the area. Registration forms and fees ($5.00, cheques payable to Animal Behavior Society Meeting) should be sub- mitted by 30 May 1977. Correspondence, requests for registration forms, and inquiries can be directed to Jon Lien, Bill Montevecchi, Cathy Noseworthy, Deane Renouf in the Department of Psychology, or John Green or Jake Rice in the Department of Biology, Memorial University of Newfoundland, St. John’s, Newfoundland AIC 5S7. water fish, reptiles, and amphibians; physiological plant ecology; population dynamics (plant, animal, and microbial); pollution and applied ecology; in- vertebrate ecology; and community ecology (plant and animal). For any further information please contact Dr. P. B. Cavers, Department of Plant Sciences, Univer- sity of Western Ontario, London, Ontario N6A 5B7 or phone 519-679-3282. The Changing Sea-bird Populations of the North Atlantic An international conference sponsored by British Ornithologists’ Union, British Trust for Ornithology, Royal Society for the Protection of Birds, Seabird Group, Scottish Ornithologists’ Club, and the Wild- fowl Trust, will be held 26-28 March 1977 at Aber- deen University, Scotland. It will enable many workers on sea-birds in Europe and on the eastern seaboard of North America to meet and find out about each other’s research. Members of the spon- soring societies, and others interested in the subject, are also encouraged to attend. Each day a topic will be covered in the morning by invited speakers. The afternoon sessions are open to papers on current research, relevant to the general theme of the conference, which may deal with more specific and specialized studies. The morning topics will be human influences, surveys, and population ecology. Members of the Aberdeen Bird Club are assisting with local arrangements. Booking forms for the conference are available from Dr. Amicia Melland, BOU Office, c/o Zoological Society of London, Regents Park, London NWI 4RY. Book Reviews ZOOLOGY Handbook of Common New Guinea Frogs By J. I. Menzies. 1976. Wau Ecology Institute, Wau, New Guinea. Wau Ecology Institute Handbook Number |. 75 pp., illus. $3.20. The large island of New Guinea has a rich and varied fauna, with over two hundred species of native frogs, some of these, however, being represented by a single or few specimens. Much more remains to be done; and undoubtedly other new species will be discovered. Dr. Menzies has chosen fifty common species, and these are described and illustrated in color in his new work. The text is broken downas follows: Introduction; A note on frog biology; The colors of frogs; The frog fauna of New Guinea—composition and origin; Further reading; A guide to the identification of frogs on the New Guinea mainland. This takes up about 16 pages. The Systematic account, which is the main body of the text, comprises some 45 pages. This is followed by a 1-page section on the preservation of frogs, a 2-page Glossary, and an important 6-page Appendix that lists the species of frogs that have been described from Papuan subregion. Dr. Menzies does Waterfowl of North America By Paul A. Johnsgard. 1975. Indiana University Press, Bloomington. 575 pp. $25. The stated purpose of this book is to provide an up- to-date account of the ecology and reproductive biology of all species of waterfowl in North America. It was with pleasure that I received it; no major work of this kind had been produced since F. H. Kort- right’s (1942) “Ducks, Geese and Swans of North America.” With the tremendous volume of data which has accumulated since 1942, an updated modern- ization of this classic work combining breeding distributions, life cycle and behavioral observations was overdue. Dr. Johnsgard has attempted to reacha large audience including naturalists, hunters, and pro- fessional biologists. This is an extremely difficult goal, and one which he has only partially attained. The introductory chapters present a useful and lucid discussion of present-day theories on the ecology of waterfowl. Interesting parallels are shown between North American and Eurasian species. Some of the material presented, however, is too simplistic to be of value. For instance, the author rightly recognizes the not (very wisely perhaps) make comment on the validity of the names mentioned in the Appendix. There is also a 2-page Index. Descriptions of the fifty species (or occasionally a species group) are of necessity brief, but should be adequate in most cases. Included in each description are also notes on variation and habitat. The color illustrations for the most part can be described as very good. For anyone interested in New Guinea frogs this work should be as useful in the library as in the field. Considering the extremely modest cost of the book one cannot possibly go wrong in purchasing a copy. Dr. Menzies is to be congratulated. It is hoped that other handbooks, now in preparation, on the flora and fauna of New Guinea, will measure up to Handbook Number |. STANLEY W. GORHAM Natural Science Department, New Brunswick Museum, Saint John, New Brunswick E2K 1E5 problems of generalizing with respect to habitats used by species of waterfowl, and then presents a summary table of habitats with its obvious draw- backs. (Not all Snow Geese nest on coastal deltas; the Banks Island colony is more than 20 km inland. Whistling Swans and Ross’ Geese are not considered high arctic breeding species and Pintails are more abundant in prairie and parkland habitats than in low arctic areas.) Many of the data presented in the tables are out of date. In some cases they are meant only to indicate trends, but kill data, Christmas counts, and monetary values are obsolete. Within these chapters a section relating particular problems facing waterfowl would have been timely and meaningful. In particular, oil pollution as it relates to arctic species and wintering populations could have been mentioned. The body of the book comprises individual species accounts including physical parameters, guides to identification, age and sex criteria, breeding and wintering distributions, biological, ecological and be- havioral data. Missing is information on migration routes, important staging areas, and times of year 103 104 when the various species are to be seen in different areas of the continent. Within the species accounts, many of the data are well-presented, particularly the summarized informa- tion on behavior. But many factual errors exist among the data on breeding distributions and in- complete discussions of habitats are presented. Examples of errors include a misquotation of Mann- ing et al. (1956), who do not state that Ross’ Geese breed on Banks Island; Snow Geese do not breed throughout Banks Island but are confined to the southwest (Manning et al. 1956); both Atlantic Brant and Pacific Brant are known to nest on Prince Patrick Island (Handley 1950); MacDonald (1960) and Godfrey (1966) report that brant breed on Ellef Ringnes Island; Gadwalls and Lesser Scaup breed in Quebec at Lake St. Francis (Ouellet 1970); no breeding records exist for Pintails on Banks Island (Manning et al. 1956; Godfrey 1966); and the range map presented for the Red-breasted Merganser is incorrect relative to evidence presented by Godfrey (1966). Most of these errors stem from an apparent incomplete review of Canadian literature on water- fowl, especially from the high Arctic, Mackenzie Valley, and Hudson Bay/ James Bay. Breeding habitats should have been dealt with ina more complete manner, specifically in the case of cosmopolitan species. No mention is made of nesting habitats of Mallard, Black Duck, Pintail, American Wigeon, or other species in boreal forest areas. Handbook of North American Birds. Volumes Edited by Ralph S. Palmer. 1976. Yale University Press, New Haven, Connecticut. 521 and 560 pp. $30 each. Waterfowl enthusiasts who collect books about their subject have long been plentiful and have been given much to collect. Now, in less than a year, those in North America have been presented with four large new volumes, by Paul Johnsgard, by Frank Bellrose and, most recently, these two stately compendia by Ralph Palmer and a team of thirty-two collaborators, including most of the best-known names in the business (about a third of them Canadian). It has taken Dr. Palmer fourteen years of hard labor and he has been triumphantly successful, particularly as some of his colleagues are almost as well known for their reluctance to write as for their knowledge of this goose or that duck. This is, happily, a book for ornithologists, rather than for duck hunters or ‘waterfowl managers.’ “The main purpose of these volumes is to present accounts of species”: thirty-five in Part 1 (whistling ducks, THE CANADIAN FIELD-NATURALIST Vol. 91 In addition to factual errors, editorial mistakes are also prevalent. The range maps inaccurately portray the Queen Elizabeth Islands; Manning et al. (1956) is noted as 1958 in the bibliography; and C. J. Lensink’s name is misspelled on page 87, to illustrate a few. This book is of value to persons who are unfamiliar with waterfowl or the published literature. But because of incorrect and incomplete data this work has not fulfilled the stated objectives and as a result does not present a true picture of the waterfowl of North America. Literature Cited Godfrey, W.E. 1966. The birds of Canada. National Museum of Canada Bulletin Number 203. 428 pp. Handley, C. O. 1950. The brant of Prince Patrick Island, Northwest Territories. Wilson Bulletin 62: 128-132. Kortright, F. H. 1942. Ducks, geese and swans of North America. Stackpole Company, Harrisburg, and Wildlife Management Institute, Washington, D.C. 476 pp. MacDonald, S. D. 1960. Report on biological investiga- tions at Isachsen, Ellef Ringnes Island, NWT. National Museum of Canada Bulletin Number 172: 90-97. Manning, T. E., E. O. Hohn, and A. H. Macpherson. 1956. The birds of Banks Island. National Museum of Canada Bulletin Number 142. 139 pp. Ouellet, H. 1970. Changes in bird fauna of the Montreal region, Canada. Canadian Field-Naturalist 84: 27-34. IAN D. THOMPSON P.O. Box 895, Cochrane, Ontario POL 1C0 2 and 3. Waterfowl, Parts 1 and 2 swans, geese, and dabbling ducks) and twenty-nine in Part 2 (diving- and sea-ducks, mergansers, wood ducks). The taxonomy is that of a conservative ‘lumper,’ i.e., mostly after that of Delacour’s Water- fowl of the World (1954-1959) trying yet again to treat the Brant as a single species, the Blue Goose as a color phase of the Snow Goose (in Anser), and des- cribing the Canada Goose by as few as eight trinomials. Eighteen of the species accounts are brief, because the birds described are not now native to North America, including the sad case of the Labrador Duck, which became extinct a century ago before much was learned about it, but these short treatments of other people’s birds keep up the high standard of the major essays, although sometimes not fully updated since first being written a decade or more ago. One of the major features of this handbook, not to be found in any recent competitor, is the very full description of all known plumages (“what the bird is 1977 wearing”). As an elderly European, reared on Schidler and Annie Meinertzhagen, I find the plumage-naming awkward, but suppose that this work will help to consolidate the use of the system introduced by Humphrey and Parkes (1959, Auk 76: 1-31). The colored figures of downy young, done originally direct from life, by Colleen H. Nelson are worthy of a book by themselves. The sections on field identification are full of acute observations. The treatments of distri- bution and migration do an effective job of boiling down what is often a welter of conflicting facts and awkward gaps into a semblance of coherence. The sections on banding status, most often little more than a record of numbers marked and recovered up to some date long past, could have been left out. The breeding biology and habits sections include many major essays and much information previously unpublished, buried in theses or agency reports or the heads of biologists. It is this “new” knowledge above all that BOOK REVIEWS 105 will make these volumes useful and enjoyable for many readers; and that will also, one hopes, prompt further studies. Although Palmer writes of the Anatidae as “already the best-known avian family” he reminds us repeatedly, if implicitly, how far we are away from an adequate view of waterfowl and their place in nature. Those of us who care for birds, even if only ducks, have many reasons to applaud the work of Palmer and his team. We have no good cause to reproach them either, though it is sad that to save space the Literature Cited section (still 27 pages in length) omits the titles of papers and gives only meagre details of books. From Abdulali (1949) to Ytreberg (1960) is a long journey. We salute the travellers; with Ralph Palmer in the van. HUGH BoYD Canadian Wildlife Service, Ottawa, Ontario KIA 0H3 Insects that Feed on Trees and Shrubs: An illustrated practical guide By Warren T. Johnson, and Howard H. Lyon, with the collaboration of C. S. Koehler, N. E. Johnson, and J. A. Weidhaas. Cornell University Press, Ithaca, New York. 464 pp. $35. The striking feature of this invaluable book is the series of 212 color plates, comprising about 1500 individual photographs, of which all but 13 were taken by the authors and collaborators. The photo- graphs depict over 700 species of pests, mostly in the living state, and their work. Explanatory text, some- times accompanied by monochrome figures, is given on the facing pages. The illustrations are not only technically and artistically excellent, but they are carefully chosen to help forester, landscaper, and gardener to recognize the pest and the damage it causes as he is most likely to encounter them. The text gives brief but useful notes on the biology and importance of each pest and often of similar species on the same or other hosts. A wide range of trees and shrub species is represented, and both eastern and western parts of the continent are dealt with. The organization of the book is logical, though the diversity of subject matter makes it a little hard to find one’s way through it. There is a main division between conifer-feeders and feeders on broad-leaved plants. Within each grouping there is a secondary classifi- cation, partly by type of damage and partly taxo- nomic, and within these categories a tertiary sub- division, again partly biological and partly taxo- nomic. A “Reader’s Aid” at the beginning gives a bird’s-eye view of this arrangement, and more detailed access is provided by the very comprehensive index. In general the emphasis is on forest and shade trees and ornamental shrubs, though there are frequent references to fruit-tree species. It would be easy to complain of pest species that are omitted: almost every reader would have his own list of neglected favorites. Actually the selection of species is very good, particularly when it is considered that it depended on the available photographic material. Though frequently suggested, the great variety of un- mentioned pests probably deserves more emphasis. Occasionally there are misleading statements in this regard, as on p. 166, “...larvae of several species of moths mine oak leaves” [there are dozens]. “There are two common species in the eastern states” [there are many]. As might be expected with so wide a coverage, there are taxonomic inaccuracies (mostly not serious), e.g., Dioryctria abietivorella Grote is listed under the old identification D. abietella (D. & S.) on p. 24. There are some misprints, e.g., on p. 222 Euzophera semifuneralis is given as semifumeralis. The treatment of authors’ names for species is erratic: they are abbreviated or not, often on the same page— Walsingham and Wlshm. on p. 36; periods are often omitted after abbreviations—Keif for Keifer on the same page; diacritical marks and accents are at least sometimes omitted from foreign names, and, though parentheses are generally used for authors’ names where appropriate, they are often omitted. 106 There is an extensive list of references. Unfor- tunately the selection is rather haphazard, and the best sources are not always cited. Dominick et al., The Moths of America North of Mexico, is not even mentioned; only one of the numerous publications by W.R. Richards on aphids and scales is listed; Chapman and Lienk on the tortricid fauna of apple in New York, Stehr and Cook on North American tent caterpillars, Freeman et al. and Morris et al. on the spruce budworm and allies, and many other key references are omitted. Probably wisely, no effort has been made to give control recommendations for the various pests. Tech- nology changes too rapidly and control measures differ too much in different environments and geographical areas for such recommendations to be appropriate in a reference work of lasting value such THE CANADIAN FIELD-NATURALIST Vol. 91 as this one. Instead the reader is advised to consult government and other information sources in his locality, and there is a useful section of such sources and how to get in touch with the agencies. Even after many years as a professional entomol- ogist and very amateur horticulturist, I found this book a revelation. It supplements ina vivid and highly informative way the standard taxonomic, agricul- tural, forestry, and horticultural manuals previously available. The authors are to be congratulated on a thoroughly successful achievement. EUGENE MUNROE Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6° The Behaviour of Ungulates in Relation to Management Edited by V. Geist and F. Walther. 1974. IUCN Publica- tions, New Series No. 24. Morges, Switzerland. 2 Volumes. 941 pp. $15. The Behaviour of Ungulates in Relation to Management is a collection of papers presented at an international symposium held at the University of Calgary, Alberta (2-5 November 1971). Contained in the two-volume text are 55 presented papers and one summary paper. The convenors had decided “that our diverse objectives could best be realized by asking senior workers to concentrate on syntheses, and by requesting those who had recently finished major field studies to report directly on their work.” Of particular interest at the symposium were papers dealing with “the relationship of ecology to social behaviour.” It would appear, however, that not all papers presented at the symposium were included. For example, reference to a presentation by A. B. Bubenik, noted in the Proceedings of the first international Reindeer and Caribou Symposium held in Alaska (9-11 August 1972), on “social well being as a special aspect of animal sociology,” appears to have been omitted. Although the conference was held in North Am- erica, only 50% of the contributors resided on this continent. The species examined, the study areas utilized, and the researchers themselves all attest to the truly international scope of this symposium. Canadians made an admirable showing either author- ing or co-authoring 12 papers. Among the general review topics in Volume One are mother-infant relationships, combat and courtship displays, and scent communication. Other papers examine specific ungulate groups such as the Equidae, the Suidae, and the African Bovidae. Volume Two presents information on the specific application of ethology to management. For instance, Laws’ paper on elephants points out the social function of the old females within elephant popula- tions. These individuals assume the leadership and infant-care responsibilities. Were these animals indis- criminately removed from the elephant population the herd’s survival would be seriously damaged. A different viewpoint is suggested by Cumming in his report on roe deer. Apparently mock fighting among young non-territorial bucks accounts for most of the tree-bark destruction caused by roe deer. The tree damage could be reduced by removing some of these young bucks without threatening the population’s ultimate survival. Another interesting example of management by utilizing ‘characteristic behavioral traits’ is Wilkinson’s description of techniques used in capturing young muskox for domestication purposes. The papers of this symposium clearly show the importance of ethology to management. Since world ungulate species represent valuable recreational re- sources and, in some cases, have significant potential as protein sources for human consumption, increased ungulate production ought to be a serious manage- ment objective. Utilization of behavioral character- istics in management practices may help meet this ob- jective. This book may help many managers and should be read. PETER CROSKERY Ontario Ministry of Natural Resources, Ignace, Ontario 1977 Mammals of the World By E.P. Walker. 1975. Third edition. Edited by J. L. Paradiso. Johns Hopkins University Press, Baltimore and London. 1500 pp. $43 for the two boxed volumes. Mammals of the World, begun by Ernest P. Walker in 1933 and first published in 1964, has proven so popular that it has recently appeared in a third edition. For those who do not know this work, it is a two-volume, attractively boxed, 1500-page compen- dium of much that is known about Recent mammals with at least one page devoted to each order, one to each family, and one to each genus. Species are often discussed specifically under genus. The text deals with such information as distribution, habitat, coloration, anatomical details, reproduction, and habits. Compared to the original volumes which have the same pagination, this 1975 edition has few changes in the text, even for mammals on which research has recently been carried out. There is no new informa- tion, for example on moose, beaver, narwhal, or Canadian deer. Apparently only if information was sent in to the Genera of Recent Mammals of the World project, sponsored latterly by the New York Zoological Society, as I sent some on the giraffe, was it incorporated into this new edition which was super- vised by John L. Paradiso after the death of Walker in 1969. In the first edition a third volume contained references to research works on mammals on which the text was based. This volume has wisely been BOTANY BOOK REVIEWS 107 omitted by Paradiso. References are soon outdated and can readily be retrieved from other sources such as Biological Abstracts and Zoological Records. One way in which this edition is superior to its predecessors is in its pictures, all black and white, which occur on almost every page. Where in the first edition there were many blank spaces, these are now often filled with photographs. For example, there are two new photos of the mule deer, while those of the pronghorn and moose have been exchanged for better pictures. Where possible, drawings have been re- placed by pictures of living animals, and there are fewer photographs of museum study skins and mounted ones to represent little-known species. Where taxonomically useful, photographs of skulls are presented. This edition of Mammals of the World contains a few anomalies. For example, why has the African tree rat of 1964, Colomys goslingi, become the African water rat of 1975? This work should be essential for every university library and for other professional libraries that havea zoological bias. The price, which 'l am sure is justified, will unfortunately prohibit many zoologists from buying it. ANNE INNIS DAGG Box 747, Waterloo, Ontario N2L 3S6 Catkin Bearing Plants (Amentiferae) of British Columbia By T. Christopher Brayshaw. 1976. British Columbia Pro- vincial Museum, Occasional Paper Number 18. British Columbia Provincial Museum, Victoria. $3.00. This book is the latest in a series of contributions to the Flora of British Columbia published by the British Columbia Provincial Museum. Dr. Brayshaw covers in his treatment the genera Populus, Salix, Myrica, Alnus, Betula, Corylus, and Quercus which are included in four families. Only two of these genera, Salix and Betula, contain a sub- stantial number of species and Salix dominates the book with 43 species. The “Amentiferae,” as the author recognizes, is not a natural group of families, but evidently they are dealt with here as a group in order to include these important families of deciduous trees and shrubs under a single cover. Salix and Betula are taxonomically very difficult plant genera and the author is to be commended for his efforts to make these plants more understandable to serious ama- teurs, wildlife biologists and botanists alike. The outstanding feature of this work is the excellent pen-and-ink illustrations done by the author himself. They are of high professional and artistic quality and aid in the recognition of the species in a way that few illustrations do. The only shortcoming of the illustra- tions is that the technique does not enable leaf pube- scence to be well displayed and Salix exigua ssp. exigua and S. commutata do not come across as the hairy plants that they are. Also it seems that the illustration of S. exigua ssp. melanopsis (Figure 22) was based on a misidentification of a specimen of ssp. interior with blunt bracts. In addition to the illustrations, detailed distribution maps are presented for each species. These maps very 108 well illustrate not only the distribution of the species in the province but also the large gaps in our knowl- edge of the flora of British Columbia. Hopefully this book will challenge collectors to collect these genera with more confidence and to learn more about these interesting plants. In some species the infraspecific taxa have been mapped using different symbols. It would have been useful if this technique had been used more often. Even in those species whose subspecies and varieties do not show any geographical distinctiveness it would be useful to make that point. In Salix barrattiana the author recognizes var. tweedyi. It would be helpful to know where this variety occurs in British Columbia, since, as far as I am aware, it does not occur in Canada. In the text each species is treated separately and a concise but adequate description is followed by brief ecological and distributional notes. Where infra- specific taxa are recognized a key is presented and sometimes these taxa are discussed. In his recogni- tion of infraspecific taxa in Salix the author seems to have accepted, or at least included, as many names as he could find. It is hard to conceive that he could have seriously studied the variation of these plants, either in the herbarium or field, and then proceeded to recognize the many varieties and forms that have been based on single characters such as leaf shape or pubescence. For example, the narrow- and broad- leaved variants of S. barrattiana, and S. planifolia; the hairy- and glabrous-leaved variants of S. bebbiana and S. sitchensis; and the capsule-pubescence variants of S. gracilis are all given formal recognition. From the point of view of a wildlife biologist or amateur botanist the use of formal nomenclature for these trivial variations must seem to complicate the matter unnecessarily. This information could have been better conveyed in a paragraph dealing with morpho- logical variation in the species. Furthermore, the use of var. hypoglauca of S. pedicellaris is difficult to justify in light of the evidence that the absence of wax on the undersurface of the leaves of the type collection is almost certainly an artifact of drying. It is generally accepted in taxonomic works for the description to be based mainly on dry herbarium specimens. Sometimes, however, this may result in misleading information for the field biologist. During a collecting trip on Vancouver Island, on which I was accompanied by the author, we tried to locate specimens of Salix geyeriana. We eventually found plants that appeared to be this species on the margins of a small lake; but the twigs were yellowish-green rather than blackish as is usually described in the literature. We discovered later that the twigs are always yellowish-green to greenish-brown in life but become black on drying. I am surprised, in view of the THE CANADIAN FIELD-NATURALIST Vol. 91 confusion that this point caused us, that Dr. Brayshaw did not attempt to clarify this for other field biologists. In most of the genera treated here, especially Populus, Betula, and Salix, hybridization is relatively common and does pose serious problems in identifi- cation of individual specimens. The author discusses the nature of hybridization and mentions that this may lead to some confusion in identification but he loses the opportunity to describe the importance of hybridization in nature and the means whereby the hybridizing species remain distinct in the face of hybrid swarms that theoretically could lead to their complete merger. In some cases I experienced some uncertainty about the author’s understanding of particular hybrids. Betula papyrifera var. subcordata is treated by the author as a variety but in the discussion he mentions that Dugle has shown it, “to be an introgressent resulting from crossing of B. papyri- fera with B. occidentalis.” If he is of the opinion that Dugle is incorrect, and that is implied by his taxonomic treatment, then he should have clearly expressed his position and reasons. In the author’s attempts to publish new taxonomic combinations or to change the taxonomic status of existing names the correct procedure according to International Rules of Botanical Nomenclature, was not followed, except in one case, and the proposed name changes are therefore not validly published. These errors are undoubtedly an oversight on the part of the author and it is regrettable that such lapses, that could have been caught by an external reviewer or a knowledgable editor, mar an otherwise creditable work. I do not wish to dwell on the negative aspects of this book, for much credit should go to the author for tackling a very difficult group of plants. The prepara- tion of keys to Sa/ix and Betula is a very difficult task indeed and the examination and identification of the many specimens and the plotting of their distribution is very time-consuming. The inclusion of taxa that are not known from the province, but to be expected there was a good bit of foresight and one of these species, Salix setchelliana, was discovered by the author even before the book went to press. There is no question that all naturalists interested in the flora of British Columbia should own this book. If it leads them, as I think it will, toa greater awareness of these important plants and helps them overcome their fears of the taxonomic problems that they present, then this work will have been more than justified. GEORGE W. ARGUS National Museum of Natural Sciences, Botany Division, Ottawa, Ontario KIA 0M8& 1977 BOOK REVIEWS 109 Atlas of United States Trees. Volume 2. Alaska Trees and Common Shrubs By Leslie A. Viereck and Elbert L. Little. 1975. Miscel- laneous Publication Number 1293, Forest Service, United States Department of Agriculture, Washington, D.C. Text 19 pp. + 105 full-page maps. Paper $3.10. This publication represents the second volume of a projected series entitled the Atlas of United States trees. The stated objective of the series is to present atlases of large-scale maps which clearly summarize the known natural distributions of native trees and shrubs. The publication of such species distribution maps has considerable value (1) in serving as a basis for many kinds of phytogeographical and related systematic or evolutionary studies, (2) for indicating to botanists where possible errors or gaps in distri- butional knowledge may exist, (3) in preserving for historical records the original natural distributional information before possible destruction or alteration of the natural vegetation by human activities, and (4) in serving such economic purposes as indicating where valuable species or possible ecotypes might be located. The authors also stress the potential value of the maps for land-use planning. Volume | of this series presented large-scale distri- bution maps of 200 conifers and 106 of the more important hardwood-tree species of the contiguous 48 states. Volume 2 contains Alaskan distribution maps (all 12 X 11” size at the scale 1:10 000) of 82 species of native woody plants including 32 trees and 50 shrubs, six of which sometimes reach tree size. Since this number includes all trees but represents hardly one half of the total number of native shrub species in Alaska, a choice was apparently made to include only the larger, more common, more widely distributed, and economically or ecologically more important shrubs. One wonders at the omission of some relatively common, or at least locally abundant, shrubs, such as numerous species of Salix, Betula occidentalis, Arctostaphylos alpina (including A. rubra), Diapensia lapponica, Rhododendron camts- chaticum, Cassiope spp. (particularly C. tetragona), Dryas spp., Phyllodoce spp., Loiseleuria procum- bens, Linnaea borealis, etc., especially when other shrubs, either just as limited in distribution, or equally as low-growing in habit, as some of these, were included. This is not intended as a serious criticism, however; the reader should note only that this volume does not constitute a complete atlas compendium of all Alaskan shrubs. Plant taxonomists will not necessarily agree with all of the authors’ concepts and treatments of the species of particular groups. In most cases, I would agree with them, although it does seem unfortunate to me that all of the tree birches have been lumped together and mapped as a single taxon, Betula papyrifera. The authors’ taxonomic treatment as well as their mapping of species distributions of the difficult willow genus, Sa/ix, is taken almost directly from Argus (The Genus. Salix in Alaska and Yukon, National Museum of Canada, Publications in Botany, Number 2, 1973), although hardly more than one third of the species recognized by the latter for Alaska are included in the present volume. Volume 2 of this Atlas essentially follows the format of Volume |, but it differs in several note- worthy respects. The county outline type of base map, which was used in Volume | for showing species’ ranges in the 48 contiguous states, is replaced by a base map of Alaska which is essentially hydro- graphic, showing rivers and lakes, on which dots or rings are superimposed to indicate the location of cities, towns, and other key places. Preceding the 82 individual species distribution maps that represent the primary objective of this volume, are 23 large-scaled general maps, showing numerous physiographic, en- vironmental, vegetational, geo-historical, political, etc., features, which are useful for a better under- standing of the species’ distributions and for allowing possible correlations to be made in their interpre- tation. The bringing together of all this geographical and especially environmental information about Alaska in these general maps should be considered one of the more valuable contributions of this publica- tion. Nevertheless the convenient transparent overlay maps by which such environmental etc. features were presented in Volume | may be missed by some readers. Unlike the maps in Volume 1, the species distribu- tions are not indicated by a black cross-hatching or gray shading of the overall ranges, but rather by dots each representing a known distributional record. The presumed overall range limits are then shown by lines drawn conservatively around the dots. The use of such dot distribution maps seems much preferable to shading simply the overall ranges, since possibly unusual “extraterritorial” occurrences are not over- emphasized. All distributional information is given in a red-brown colour and thus is clearly set off from the clutter of the black on white base-map features. The species ranges have not been mapped outside of Alaska. The text, however, contains a brief paragraph for each included species which summarizes both the Alaskan and extralimital ranges. For more ample species descriptions, botanical drawings, and ecologi- cal information, the reader is referred to Viereck and Little (Alaska trees and shrubs, Agriculture Hand- book Number 410, Forest Service, United States Department of Agriculture, Washington, D.C., 1972, $3.25). 110 The species maps show known presence or absence, but not abundance or density; thus, these maps are not meant to indicate forest cover or vegetational types. This is an important distinction which would seem to undermine somewhat the authors’ repeated conten- tion that these distribution maps should serve as a valuable basis for land-use planning. As in Volume 1 of this atlas series, an attempt has been made to record only the known, presumably natural occurrence of each species, but not to explain or speculate on how or why it occurs where it does; such interpretations are left for other phytogeographers to make using the information made available here. The users of atlases such as Volumes | and 2 of the present series will be concerned about the reliability of the maps. The species maps of Volume 2 may well be more reliable, although likely less complete, than those of Volume 1, largely because they have been compiled from fewer and these quite reliable literature sources, which have then been amplified by herbarium specimens mostly seen by the authors and sight records that are also mostly those of the authors themselves. The primary original basis for most of ENVIRONMENT Ecological Sites in Northern Canada Edited by David N. Nettleship and Pauline A. Smith. 1975. Canadian Committee for the International Bio- logical Programme Conservation Terrestrial—Panel 9. Canadian Wildlife Service, Ottawa. 330 pp. Paper $3.75. There are areas in northern Canada of biological, geological, and historic importance which urgently require special attention. The scope and intensity of human activity is expanding rapidly in Canada’s north and pristine wilderness areas are now threat- ened. This is the subject matter of the publication. In 1968, the Canadian Committee for the Inter- national Biological Programme (IBP) designated panels for 10 geographical regions in Canada. Panel 9, Arctic tundra areas, and Panel 10, Subarctic areas, fall within the Yukon and Northwest Territories. Panel 9 sites are described in this publication. The objectives of the northern panels were these: (1) to locate and describe representative examples of natural arctic and subarctic ecosystems in co- operation with local residents, industry, and the Federal, Northwest, and Yukon Territorial Govern- ments; (2) to demonstrate how the biological values of each potential site may equal or outweigh all other values of that site; and THE CANADIAN FIELD-NATURALIST Vol. 91 these species distribution maps appears to have been Hultén (Flora of Alaska and neighboring territories, Stanford University Press, California, 1968), or inthe case of Salix species, Argus (1973). The locality records for the species maps might well have been considerably amplified, although perhaps not signifi- cantly enough to change many range limit lines, if the authors had checked more specimens available in various unconsulted herbaria which contain sizable Alaskan collections. This volume is recommended to anyone who has an interest in the distribution of native trees and shrubs of Alaska and adjacent Canada. It would seem especially useful for botanists (systematists, ecolo- gists, phytogeographers, evolutionists, etc.), foresters, and naturalists if used in conjunction with the 1972 book, Alaska trees and shrubs by the same authors. VERNON L. HARMS Fraser Herbarium, Department of Plant Ecology, University of Saskatchewan, Saskatoon S7N 0WO (3) to aid the three governments in providing for the preservation of these biologically important areas in the form of Ecological Sites. Ecological Sites are designated special areas en- compassing a variety of plant and animal communi- ties. Many of these communities contain relict or endangered populations, unique plant associations, breeding areas, and critical habitat for wildlife. All interested parties cooperated to ensure that the traditional hunting and fishing privileges of indig- enous people were protected. Also, necessary and realistic consideration was given to gas, mineral, and oil development activities. Many proposed sites can support multiple land use, while other sites of high biological value may be too sensitive to disturbance. Panel 9 is divided into seven geographical regions. A total of 71 proposed sites is discussed in the publication. Each site is identified by geographical name, plus latitude and longitude. Each entry is accompanied by a map of an appropriate scale. Geographical and topographical features of each site are discussed, plus exceptional features such as bird, mammal, and marine popula- tions, migration routes, critical wildlife habitat, vegetative patterns, archeological sites, etc. 1977 The extensive bibliographic references relating to each site are perhaps the most valuable aspect of this volume. Literally hundreds of bibliographic entries are listed and enthusiastic northern students can derive immense knowledge by pursuing these ref- erences. One important aspect of each entry is an indication of the protective status. Of the 71 sites discussed, only 12 receive some form of official protection as portions of Migratory Bird Sanctuaries, Game Sanctuaries, or National Parks. Therefore, the casual reader must not infer that the sites listed are “saved,” as it were. These sites are proposals only and will be subject to review before receiving official IBP status. The reviewer has been fortunate enough to have viewed several of these areas personally. Areas such as Environmental Change in the Maritimes Edited by J. G. Ogden, III and M. J. Harvey. 1975. Nova Scotian Institute of Science, Halifax, 109 pp. $5.00. This little book, published as a supplement to the proceedings of the institute, is the result of a symposium in Quaternary history held at Dalhousie University in 1971. Change began, for purposes of the symposium, as melting glacial ice began to uncover Nova Scotia and New Brunswick about 13 000 years ago. The environment, studied largely for its own sake, is presumably defined as the human environ- ment. It is the geography of land and sea, of the vegetation and fauna, terrestrial and marine. Change in the physical environment of organisms, change in their distribution through ecology and migration, are both studied, and change in the human habitat, reflecting in part man’s impact on nature, is studied for the prehistoric and recent past. The writing varies as in most symposia from the quite simple and popular to that which makes few concessions to the non-scientist. All have made an effort to adjust to general communication, simplify- ing with maps and tables, if not in the language. It isa praiseworthy attempt to set forth at an early date the results of recent work on geology and biology of the Maritime Quaternary. But, although one contributor has made an effort to update his bibliography, it is regrettable that four years should have elapsed between symposium and publication. Ogden introduces the symposium with the idea that quite a small change in such large systems as climate may bring about major change in biology. Even glaciation may be a “small change” amplified. But Terasmae details sixteen categories of hypotheses of climatic change; evidently, it is not to be explained BOOK REVIEWS 111 Padle-Kingnait Fiord, Baffin Island; Cape Searle, Baffin Island; Anderson River, Region 4 and Caribou Hills, Mackenzie River Delta are all worthy of IBP status with high biological and aesthetic value. This book contains very worthwhile information for developers, development-oriented agencies, and northern planners in general. The bibliographic references are a real plus and make the book a valuable tool for anyone interested in northern con- servation, ecology, geology, and archeology. This volume belongs on any Arctic reference bookshelf. DAN MURPHY Fish and Wildlife Service, Government of the Northwest Territories, Yellowknife, Northwest Territories XOE 1HO simply. He uses the Bryson-Borchert hypothesis of the correspondence of vegetation zones with the domi- nance by a particular combination of winter and summer air masses, to emphasize the relationship of vegetation and climate and to pose the general questions. Railton and Mott report on postglacial vegetation change as exhibited by pollen sequences, the former from Nova Scotia, the latter from New Brunswick. The only symmetrical arrangement of radiocarbon dates suggests to the former a local ice-cap centered about Kejimkujik National Park and persistent till nearly 7620 BP. The latter finds retreat of the ice margin across New Brunswick extremely rapid. In Railton’s article, many divergences in interpretation between his findings and Livingstone’s are empha- sized. The pollen record is not as yet fully harmonized. Faunistic studies show an important effect on littoral shellfish (Bousfield and Thomas) of changes in sea temperature, and strong effects on the distribution of beetles—and, by extension, other insects?—from human economic activity (Howden). Byers shows the effects of prehistoric man on fauna: Pleistocene over- kill is more likely a result of indifference and omnivory than of overspecialization on a favorite game (mastodon). Fire, he suggests, was introduced into this susceptible forest rather to modify habitat for moose hunting than for its very limited agriculture. Mann illustrates the impact of recent economic demands on the environment in referring to the endangering of the Atlantic salmon population and the pollution of Maritime shores. Perhaps our comfort as to the future must be sought in such technological feats as now maintain salmon on the St. John River above Mactaquac Dam. 112 Grant reviews the evidence as to rates of coastal submergence, its location and causes, thus providing a most interesting supplement to Goldthwait’s classic of 1924, The Physiography of Nova Scotia. In con- clusion, Ogden suggests the value of certain isotopes as measures of change, such as pollution, in the environment. Has the Quaternary in general not seen rapid change in the biota of the Maritimes and its environ- ment? Should man’s activities be looked upon as a mere fluctuation in the geological record, another NEW TITLES Zoology Advances in the study of behavior. Edited by J. S. Rosen- blatt, R. A. Hinde, E. Shaw, and C. Beer. Volume 6. Academic, New York. 304 pp. $19.50. + America’s master of bee culture. The life of L. L. Langs- troth. 1976. By Florence Naile. Cornell University Press, Ithaca, N.Y. 215 pp. $9.95. Amphibians and reptiles of the Lake Michigan Drainage Basin. 1976. By Edwin D. Pentecost and Richard C. Vogt. Volume 16, Environmental status of Lake Michigan region. Argonne National Laboratory, Argonne, Ill. (Order from U.S. National Technical Information Service, Springfield, Va.) 69 pp. Paper $4.50. * Anatomy of fishes, part I: text and part II: figures and plates. 1976. By Wilhelm Harder. Translated by Stephen Sokoloff. 2nd edition. Schweizerbart’sche, Stuttgart. xii+ 612 pp. and ii+ 132 pp., illus. DM 238 ($96.40). Animal communication by pheromones. 1976. By H. H. Shorey. Academic, New York. 176 pp. $16.50. Aspects of sponge biology. 1976. Edited by F. W. Harrison and R. R. Cowden. Academic, New York. 371 pp. $16.50. | Biology of the Kaminuriak population of barren-ground caribou. Part 4: Growth reproduction and energy reserves. 1976. By T. C. Dauphiné, Jr. Canadian Wildlife Service, Ottawa. Report Series No. 38. 69 pp., illus. Paper $3.25 in Canada, $3.90 elsewhere. + Bird hazards to aircraft. 1976. By H. Blokpoel. Clarke Irwin, Toronto. xiv + 236 pp., illus. $9.50. Bird sounds. 1976. By Gerhard A. Thielcke. Translated from German edition (1970). University of Michigan Press, Ann Arbor. viii + 190 pp., illus. Cloth $6.95; paper $2.95. + The birds of New Brunswick. 1976. By W. Austin Squires. 2nd edition. New Brunswick Museum, Saint John. Mono- graphic Series No. 7. v + 220 pp, illus. THE CANADIAN FIELD-NATURALIST * * Vol. 91 change among many? Since the ice began to melt away, man seems to have induced a series of ever more rapid changes. May not change in the freshwater and coastal environments be as great as the changes brought about in flora, insect, and other terrestrial faunas-over the past two centuries? DAVID ERSKINE 71 Green Bush Road, Willowdale, Ontario Checklist of the world’s birds. A complete list of species, with names, authorities and areas of distribution. 1976. By Edward S. Gruson. Quadrangle (New York Times), New York. xii + 212 pp. $10.95. Environmental toxicity of aquatic radionuclides. Models and mechanisms. 1976. Edited by M. W. Miller and J. N. Stannard. Ann Arbor Science, Ann Arbor. xv + 333 pp. $26.50. Field guide to Pacific fish. 1976. By David Somerton and Craig Murray. Douglas, Vancouver. 96 pp., illus. $4.95. Fishes of the world. 1976. By JosephS. Nelson. Wiley, New York. 416 pp., illus. $24. Physiology of the Amphibia. 1976. By Brian Lofts. Academic, New York. Volume 3. 520 pp. $58.50. + Policy for Canada’s commercial fisheries. 1976. By anony- mous. Environment Canada, Fisheries and Marine Service, Ottawa. 70 pp. + appendices. Free. Botany + Illustrated flora of Illinois. Sedges: Cyperus to Scleria. 1976. By Robert H. Mohlenbrock. Southern Illinois University Press, Carbondale. xii + 192 pp., illus. $15. Vascular plants of British Columbia. A descriptive resource inventory. 1977. By Roy L. Taylor and Bruce MacBryde. University of British Columbia Press, Vancouver. 772 pp. Paper approx. $28. Flora Europaea. Volume 4. Plantaginaceae to Compositae (and Rubiaceae). 1976. Edited by T.G. Tutin, V. H. Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S.M. Walters, and D. A. Webb. Cambridge University Press, Cambridge. * The chemotaxonomy of plants. 1976. By Philip M. Smith. Edward Arnold, London (Canadian distributer Macmillan, Toronto). 1977 The mosses of southern Australia. 1976. By George A. M. Scott and Ilma G. Stone. Illustrated by Celia Rosser. Academic, New York. 514 pp. $46. * Spring flora of Wisconsin. 1976. By Norman C. Fassett. 4th edition revised and enlargened by Olive S. Thomson. University of Wisconsin Press, Madison. 413 pp., illus. $8.50. * Rocky mountain flora. 1976. By William A. Weber. 5th edition. Colorado Associated University Press, Boulder. 477 pp., illus. Text edition $9.50; trade edition $12.50. t Modern methods in forest genetics. 1976. Edited by J. P. Miksche. Springer Verlag, New York. xvi + 288 pp. $23.80. The seeds of dicotyledons. 1976. By E. J. H. Corner. Cam- bridge University Press, New York. Two volumes. Vol. 1: xii + 312 pp. $39.50, and vol. 2: viii + 552 pp., illus. $65. Environment } Structure and dynamics of land use. Ecologie de la zone de lAéroport International de Montréal. 1975. By Peter Brooke Clibbon. Les Presses de l'Université de Montréal. Xx1X + 369 pp., illus. $12. { The land that never melts. Auyuittuq National Park. 1976. Edited by Roger Wilson. Peter Martin, Toronto. x + 212 pp., illus + map. Cloth $15; paper $5.95. An introduction to biological rhythms. 1976. By John D. Palmer. Academic, New York. 392 pp. $19.50. + Ecology and management of animal resources. Ecologie de la zone de ’Aéroport International de Montréal. 1976. By BOOK REVIEWS 113 J. Roger Bider, Eric Thompson, and R. W. Stewart. Les Presses de l'Université de Montréal. xxii + 246 pp., illus. $12. The natural history of New Zealand. An ecological survey. 1973. Edited by G.R. William, A. H. Reed and W. W. Reed. Tuttle, Vermont. 434 pp., illus. $27.95. + Nature quizzes for Canadians. 1976. By Vicky and Bill McMillan. Douglas, Vancouver. 144 pp., illus. Paper $3.95. + Le cadre d’une recherche écologique interdisciplinaire. Ecologie de la zone de ? Aéroport International de Montréal. 1976. By Pierre Dansereau. Les Presses de l'Université de Montréal. 343 pp., illus. $12. + Plants and animals of the Pacific northwest. An illustrated guide to the natural history of western Oregon, Washington, and British Columbia. 1976. By Eugene N. Kozloff. Douglas, Vancouver. ix + 264 pp., illus. $17.50. Other + Athlon. Essays on paleontology in honour of Loris Shano Russell. 1976. Edited by C. S. Churcher. Royal Ontario Museum, Toronto. 286 pp., illus. $15. The uncertain future of the lower Fraser. 1976. Edited by Anthony H.J. Dorcey. University of British Columbia Press, Vancouver. 200 pp. Paper $4.95. Water publications of State agencies—first supplement. 1976. Edited by G.J. Giefer and D.K. Todd. Water Information Center, Huntington, New York. $23. tAvailable for review *Assigned for review THE OTTAWA FIELD-NATURALISTS’ CLUB BY-LAWS (Approved by Council 15 November 1976) IF 10. General Meeting The Club shall hold at least two general meetings each year at which the affairs of the Club shall be discussed. One such meeting, the Annual Business Meeting, shall be for the purpose of electing officers and additional members of the Council and for conducting such other business as shall arise. . Fiscal Year The fiscal year of the Club shall be the calendar year. . Duties of the Membership Committee The Membership Committee shall keep the Council informed of the state of the membership. It shall ensure that accurate records of memberships are maintained. It shall act as a liaison between the members and the Council on all matters respecting the conditions of membership. It shall recommend to the Council candidates for Honorary Membership. . Duties of the Finance Committee At the commencement of each fiscal year the Finance Committee shall prepare for the Council’s approval a statement of estimated revenues and a proposed allocation of funds for the year. It shall act in an advisory capacity to the Council in all matters concerning investments and disbursements of funds, and any other financial dealings and transactions of the Club. . Duties of the Excursions and Lectures Committee The Excursions and Lectures Committee shall make arrangements for excursions, lectures, and other activities for the education and entertainment of members. . Duties of the Publications Committee The Publications Committee shall supervise the policy, finances, and distribution of the Club’s publications. It shall act in an advisory capacity to the Council in all matters pertaining to publications of the Club. The Publications Committee shall recommend Editors of each publication for approval by the Council. It shall appoint Associate Editors for each publication. . Duties of the Nominating Committee A Nominating Committee, consisting of three members, shall be chosen by the Council early in the year. The President shall not be a member of the Committee and no Officer of the Club shall be chairman. The Committee will prepare a slate of Officers and candidates to be elected to the Council at the following Annual Business Meeting. It shall be the responsibility of the Committee to ensure that all members of the Club, whatever their place of residence, have a reasonable opportunity to make nominations. All nominations made to the Committee shall be made in writing, and shall include a statement from the nominee that he or she is willing to serve. No nominations will be received at the Annual Business Meeting, and the Nominating Committee shall ensure the presentation of sufficient candidates to satisfy the requirements of the Council as.laid down in the Constitution. . Special Committees Special Committees may be formed by the Council and delegated authority for specific tasks. Each Special Committee shall include at least one member of the Council who need not be chairman of the Committee. . Duties of the Editors The Editor of each publication of the Club shall be responsible for the editorial policy, content, and preparation of that publication. Each Editor shall be a member of the Publications Committee and shall be responsible to that Committee. The Associate Editor(s) of each publication shall assist the Editor in the preparation of that publication. Business Managers A Business Manager for The Canadian Field- Naturalist shall be appointed as specified by the Constitution. In addition, the Council may appoint a Business Manager for any other publication or for the Club itself. The duties of the Business Managers shall be as specified by the Council. 114 1977 By-LAWS MS) 11. 14. 16. 18. Treasurer’s Assistant The Council may appoint a Treasurer’s Assistant who shall be responsible solely to the Treasurer. The Treasurer will define the duties of the Assistant in consultation with the Business Manager(s) and the Chairmen of the Standing Committees. . Disbursements of Club Moneys Disbursements of Club moneys shall be made by the Treasurer on receipt of properly rendered accounts verified by the Chairman of the Committee concerned, or by a Business Manager, or as specified by the Council. Disbursements of Club moneys shall be made only by cheque bearing the signatures of any two of the three following members of the Council: President, Treasurer, Business Manager of The Canadian Field- Naturalist. . Order and Conduct of Business at Meetings The order of business at the Annual Business Meeting and meetings of the Council shall be: . Minutes of the previous meeting . Business arising out of the minutes . Communications . Treasurer’s report . Reports of Committees . New business The order of business may be changed by a unanimous vote of members present at such meetings. All above meetings shall be conducted according to the Constitution, By-laws, special rules of the Club, and normal parliamentary procedure. In all cases where differences of opinion arise, Bourinot’s Rules of Parliamentary Procedure shall be followed. Voting by Proxy will not be permitted at meetings of the Council. Annual Reports The Chairman of each Standing and Special Committee shall submit a report of the Committee’s activities during the year to a meeting of the Council. The Recording Secretary shall prepare an Annual Report at the end of the fiscal year which shall include the minutes of the previous Annual Business Meeting and accounts of the activities of each Committee. The Annual Report shall be presented at the Annual Business Meeting and shall be published in The Canadian Field- Naturalist. NnNhWN . Annual Dues The schedule of annual dues shall be as follows: Memberships — Individual: $7.00 Family: $9.00 Subscription Fees The schedule of subscription fees shall be as follows: The Canadian Field-Naturalist — Individual: $ 7.00 Libraries and Institutions: $15.00 Trail & Landscape — Libraries and Institutions: $7.00 . Expulsion from the Club Any individual may be expelled from the Club for conduct or activities prejudicial to the objectives and well-being of the Club, by a two-thirds majority vote of the elected Council, the individual first having had an opportunity to defend himself before the Council prior to the vote being taken. Amendments An amendment to these By-laws may be adopted at any meeting of the Council, by a two-thirds majority of the members present, due notice embodying a copy of the proposed amendment having been given at a previous meeting of the Council. Any such amendment shall be published in an early issue of The Canadian Field- Naturalist. Instructions to Contributors Content The Canadian Field-Naturalist is a medium for publica- tion of scientific research papers in all fields of natural history. As the journal has a flexible publication policy, items not covered in the traditional sections (Articles, Notes, Letters, News and Comment, and Book Reviews) can be given a special place provided they are judged suitable. Naturalists are also encouraged to support local natural history publications. Manuscripts Please submit, in either English or French, three complete manuscripts written in the journal style. The research reported should be original. It is recommended that authors ask qualified persons to appraise the paper before it is submitted. Also authors are expected to have complied with all pertinent legislation regarding the study, disturbance, or collection of animals, plants, or minerals. Type the manuscript on standard-size paper, if possible use paper with numbered lines, double-space throughout, leave generous margins to allow for copy marking, and number each page. Generally words should not be abbre- viated but use SI symbols for units of measure. Under- line only words meant to appear in italics. If only one or two references are cited, they can be inserted into the text. Follow the literature cited with the captions for figures (numbered in arabic numerals and typed together on a separate page) and the tables (each titled, numbered consecutively in arabic numerals, and placed on a separate page). Mark in the margin of the text the places for the figures and tables. For articles, provide arunning head, a bibliographic strip, and an abstract. Extensive tabular or other supplementary material not essential to the text should be submitted in duplicate on letter size paper for the Editor to place in the Depository of Unpublished Data, CISTI, National Research Council of Canada, Ottawa, Canada KIA 0S2. A notation in the published text should state that the material is available, ata nominal charge, from the Depository. The CBE Style Manual, 3rd edition (1972) published by the American Institute of Biological Sciences, is recom- mended as a guide to contributors. Webster’s New Inter- national Dictionary and le Grand Larouse Encyclopédique are the authorities for spelling. Tllustrations—Photographs should have a glossy finish and show sharp contrasts. Photographic reproductions of line drawings, no larger than a standard page, are preferable to large originals. Prepare line drawings with India ink on good quality paper and letter (not type) descriptive matter. Write author’s name, title of paper, and figure number on the lower left corner or on the back of the illustration. Two copies of figures in addition to originals are required for use by referees. Special Charges Authors must share in the cost of publication by paying $40.00 for each page in excess of six journal pages, plus $5.00 for each illustration (any size up to a full page), and up to $40.00 per page for tables (depending on size). Reproduc- tion of color photos is extremely expensive; price quotations may be obtained from the Business Manager. When galley proofs are sent to authors, the journal will solicit on a voluntary basis a commitment, especially if grant or institutional funds are available, to pay $40.00 per page for all published pages. Authors may also be charged for their changes in proofs. Limited journal funds are available to help offset publication charges to authors with minimal financial resources. Requests for financial assistance should be made to the Editor when the manuscript is submitted. Reprints An order form for the purchase of reprints will accompany the galley proofs sent to the authors. Reviewing Policy of The Canadian Field-Naturalist Manuscripts submitted to The Canadian Field-Naturalist are normally sent for evaluation to an Associate Editor and at least one other reviewer. Authors are encouraged to suggest names of suitable referees. Reviewers are asked to give a general appraisal of the manuscript followed by specific comments and recommendations. Almost all manu- scripts accepted for publication have undergone revision— sometimes extensive revision and reappraisal. The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain scientific quality and overall high standards of the journal. TABLE OF CONTENTS (concluded) Genetic variants in Canada of the rainbow trout, Sa/mo gairdneri, called golden trout and palomino trout ROBIN E. CRAIG and E. J. CROSSMAN Trauma-induced paralysis in a moose calf GORDON A. CHALMERS and MORLEY W. BARRETT A recent record of meadow jumping mouse, Zapus hudsonius, in the Northwest Territories RICHARD J. DOUGLASS and ANDREW E. L. McNAUGHTON Swarming of dragonflies noted at Drag Lake, Ontario T. E. PERRY, M.S. PERRY, and J. E: K. PERRY Two recent bobcat (Lynx rufus) specimens from southern Ontario DAVID NAGORSEN and RANDOLPH L. PETERSON News and Comment Book Reviews Zoology: Handbook of common New Guinea frogs — Waterfowl of North America — Handbook of North American birds. Volumes 2 and 3. Waterfowl Parts 1 and 2 — Insects that feed on trees and shrubs: an illustrated practical guide — The behaviour of ungulates in relation to management — Mammals of the World Botany: Catkin bearing plants (Amentiferae) of British Columbia — Atlas of United States Trees. Volume 2. Alaska trees and common shrubs Environment: Ecological sites in northern Canada — Environmental change in the Maritimes New Titles Mailing date of previous issue 28 January 1977 The Ottawa Field-Naturalists’ Club By-laws Corrigendum 93 94 1972. Canadian Field-Naturalist 86(3): 294. Thayer’s Gulls wintering off western Newfoundland by R. G. B. Brown. At the time this note was published a lot less was known of Thayer’s Gull (including the range) than is known now. Dr. Brown has already published a correction in the At/as of eastern Canadian seabirds. This reads as follows: “Several winter identifications off western Newfoundland (Brown 1972b) are now thought to be doubtful.” Although there is always a certain risk in publishing such sight records, even those made by competent professionals, we feel that, in total, the usefulness of these accounts greatly outweighs any harm. Certainly anyone making a compilation of sight records has to exercise judgment. THE CANADIAN FIELD-NATURALIST Volume 91, Number 1 Articles Distribution and abundance of waterfowl wintering in southern Quebec AUSTIN REED and ANDRE BOURGET Morphology, reproduction, diet, and behavior of the arctic hare (Lepus arcticus monstrabilis) on Axel Heiberg Island, Northwest Territories GERALD R. PARKER Life history observations of three species of snakes in Manitoba PATRICK T. GREGORY Status and habits of the cougar in Manitoba ROBERT W. NERO and ROBERT E. WRIGLEY Changes in small mammal populations after clearcutting of northern Ontario black spruce forest ARTHUR M. MARTELL and ANDREW RADVANYI Morphological parameters and spring activities in a central Ontario population of midland painted turtle, Chrysemys picta marginata (Agassiz) T. H. WHILLANS and E. J. CROSSMAN Germination requirements of Alaskan Rosa acicularis R. DENSMORE and J. C. ZASADA Nesting biology of the Sora at Vermilion, Alberta JAMES K. LOWTHER Notes Predation by wolves on wolverines BRUCE K. BOLES Chipping Sparrow hanged E. R. FILLMORE and R. D. TITMAN Sources of mortality in concentrated garter snake populations MICHAEL ALEKSIUK Micro-habitat selection of chestnut-cheeked voles (Microtus xanthognathus) RICHARD J. DOUGLASS and KRISTIN S. DOUGLASS Variation in selected meristic series in the golden shiner, Notemigonus crysoleucas (Mitchill), in the New Brunswick — Nova Scotia Border Region J. W. CARPENTER and C. G. PATERSON Unreliability of strip aerial surveys for estimating numbers of wolves on western Queen Elizabeth Islands, Northwest Territories FRANK L. MILLER and RICHARD H. RUSSELL The flowering phenology of common vascular plants at Bailey Point, Melville Island, Northwest Territories GERALD R. PARKER Gestation, litter size, and number of litters of the red squirrel (Tamiasciurus hudsonicus) in Quebec JEAN FERRON and JACQUES PRESCOTT The fern Woodsia obtusa (Spreng.) Torrey in Ontario DONALD M. BRITTON First report of the tiger trout hybrid, Sa/mo trutta Linnaeus X Salvelinus fontinalis (Mitchill), in Alberta JAMES H. ALLAN The Cattle Egret in British Columbia R. WAYNE CAMPBELL and WAYNE C. WEBER A Great Blue Heron preying on shiner perch in deep water JEAN-Guy J. GODIN Spotted Redshank sighted in southern Ontario H. H. AXTELL, P. BENHAM, and J. E. BLACK Fieldfare in Ontario DAVID J. T. HUSSELL and MICHAEL J. PORTER 1977 19 28 41 47 58 63 68 69 70 72 74 77 81 83 84 85 87 88 90 9] concluded on inside back cover ISSN 0008-3550 The CANADIAN FIELD-NATURALIST 1 Published by THE CTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Ganadau Volume 91, Number 2 April-June 1977 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Their Excellencies the Governor General and Madame Jules Léger The objectives of this Club shall be to promote the appreciation, preservation, and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and co-operate with organizations engaged in preserving, maintaining, or restoring environments of high quality for living things. Members of Council* President: R. A. Foxall E. Beaubien J. Murray : , W. J. Cody M. Ney Vice President: R. Taylor jl. Dicanen Gl Nicholson Recording Secretary: D. R. Laubitz A. Dugal G. Oyen Corresponding Secretary: S. Armstrong be dl BSE Go Fatenapde ; asi C. Gruchy J. K. Strang Treasurer: P. J. Sims J. E. Harrison S. M. Teeple Past President: E. C. D. Todd B. Henson C. G. van Zyll de Jong H. N. MacKenzie * This Council is in office until the Annual Business Meeting in January 1978. Correspondence: Address to The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada K1Y 4J5 The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club with the assistance of contributions from the National Research Council of Canada and The Canadian National Sportsmen’s Show. Opinions and ideas expressed in this journal are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Editor: Lorraine C. Smith Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy Associate Editors C. D. Bird Charles Jonkel David P. Scott E. L. Bousfield J. Anthony Keith Stephen M. Smith Francis R. Cook Charles J. Krebs Robert E. Wrigley A.J. Erskine George H. La Roi Copy Editor: Marilyn D. Dadswell Business Manager: W. J. Cody Production Manager: Pauline A. Smith Box 3264, Postal Station C Chairman, Publications Committee: C. G. van Zyll de Jong Ottawa, Canada K1Y 4J5 Subscriptions and Membership Subscription rates for individuals are $7.00 per calendar year. Libraries and other institutions may subscribe at the rate of $15.00 per year (volume). The annual membership fee of $7.00 includes club publications. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada K1Y 4J5. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Back Numbers Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field-Naturalists’ Club, 1879-1886, and The Ottawa Naturalist, 1887-1919, may be purchased from the Business Manager. All material intended for publication should be addressed to the Editor: Dr. Lorraine C. Smith, Department of Biology, Carleton University, Ottawa, Ontario, Canada KIS 5B6 Cover: A vibrating tail (blurred) and characteristic striking coil warn that this Prairie Rattlesnake, encountered as it was making a midday crossing of a gravel road northwest of Medicine Hat, Alberta, should not be molested. Photo by George B. Pendlebury. See article page 122. The Canadian Field-Naturalist Volume 91, Number 2 April-June 1977 Editorial Policy From time to time it is useful to state current editorial policy; not only does this remind or clarify to authors and readers our ongoing practices, but it also serves to introduce new, updated, or flexible policies. Journal editors normally endeavor to improve and standardize their publications. The Canadian Field- Naturalist, a medium for publication of original scientific research papers and notes on Canada’s natural history, has earned a well-deserved reputation among scientists and naturalists; it is internationally recognized, listed in Current Contents, and financially supported by the National Research Council of Canada. It is an outlet for the work not only of scientists but also of well- informed amateur naturalists; that is, for all people who have significant contributions to make to our understanding of many aspects of natural history. The journal, of course, takes no responsibility for statements made by contributors. Our aim is to keep the quality and overall standards of the journal as high as possible while continuing to serve our authors and readers. It is the Editor’s responsibility to ensure that the journal continues to fulfil its present important role of recording, interpreting, and communicating information on Canadian natural history phenomena. Because we are striving to improve the standards of scientific writing so it is clear, well organized, concise, simple, logical, and interesting, we often help authors to rewrite, sometimes to reorganize, to prune, and generally to improve their manuscripts. By doing this we are often able to salvage and eventually publish papers that we would otherwise reject; at the same time we are upholding the standards of the journal. To communicate their work properly, authors need to take into account the requirements of the average reader; it is especially important for our authors not to isolate themselves in their disciplines. Because many journals now cater to specialists, we hope that The Canadian Field- Naturalist, with its broad coverage of subjects and its many-sided approaches to natural history, will help to integrate studies and have a generalizing influence in the world that is becoming increasingly specialized. Although specialized papers do have a place in the journal, it is best to remember that they become more important if they are interpreted for readers other than specialists in the field of the paper. Moreover, if their message is interesting enough to be recorded in the primary scientific literature, then it should be conveyed clearly and as concisely as possible. As well, authors owe it to themselves to present their data in the best possible way. Perhaps the most common comments an editor makes are “spell out more clearly” and “rewrite more succinctly.” But we often have to tell authors, “Focus on the stated objectives and tell us the advance in science and its significance, put your contribution into perspective with what is known about the subject, delete ancillary material and parts that are redundant, and make your writing more direct and less complex.” For simplicity and clarity authors are also encouraged to use the first person and the active voice. We will not publish manuscripts that are scientifically unsound, are unoriginal, lack sufficient worthwhile new information, have been published elsewhere, or are not relevant. Preliminary studies or piecemeal submissions are also generally not acceptable. Referees are encouraged to make constructive comments basing their appraisals on sound scientific knowledge and experience and on the significance of the contribution to the ever-expanding literature. If they choose to do so, and most do, they may remain anonymous. I am convinced that the present referee system, although not perfect, is both good and essential and especially so for our 117, 118 THE CANADIAN FIELD-NATURALIST Vol. 91 journal, as the Editor cannot be cognizant with all fields of natural history. Broadminded authors, however, do make the Editor’s work easier. Often opinions are divergent and the Editor must make the final decision whether a paper is acceptable for publication. Although authors are always free to rebut the criticisms of referees and the decision of the Editor, their refutations should be based on sound arguments. Content We are, of course, aware that not everyone agrees with our editorial policies. Therefore, on occasion, we try to take a fresh look at the appropriateness of the manuscripts we publish. For example, although some journals are no longer accommodating descriptive studies and similar papers, The Canadian Field-Naturalist will continue to accept these if they contain data basic to the advancement of science. There is, however, no need for the journal to publish most management or laboratory studies as other outlets are more appropriate. But the journal does not limit itself to field studies, because laboratory, museum, and other studies may have considerable significance to the field situation. Thus the subjects of all manuscripts are given objective consideration. The journal, however, must remain true to its original purpose, that is, to give primary consideration to manuscripts that help elucidate and have some basic relevance for Canadian natural history. Studies done outside Canada are eligible for submission provided they meet this requirement for content; for example, ecosystems and/or biotas in the northern United States are often the same as those in Canada. Should we continue to publish range extensions? Last year Associate Editor David P. Scott wrote, “Tam more and more concerned with the amount of space which is being devoted to range extensions and/or speculative zoogeography. I wonder if you could consider the idea of having a single annotated list of new ranges published once a year rather than a series of notes and papers as is presently the case?” I solicited the opinions of all our other Associate Editors as well as other interested and concerned persons and received widely divergent views. An editorial by Dave Scott will be published in an early issue of the journal to elaborate on his opinion and to comment on other views. Certainly there are advantages to having a much-reduced and standardized style, but the topic is an important one and no decision will be taken without carefully weighing the pros and cons. For the time being, therefore, the current method of reporting range extensions will be continued; the merit of each paper and whether the range extension is significant will be judged individually by competent referees. Occasionally our practice of publishing sight records (that is, records not supported by either a specimen or a photograph) is questioned. Of course, there is always a certain risk in publishing sight records, even those by competent professionals, but we feel that, in total, the usefulness of these accounts outweighs their harm. Generally the species concerned are reasonably easy to identify and experienced observers clearly detect the diagnostic characteristics. Certainly anyone compiling and analyzing sight records has to appraise each one carefully. Another contentious subject is “lists.” In general we would like to discourage submission of long annotated lists of species with detailed information about each. This does not imply that we consider such lists as unimportant or of too little significance to be recorded but, based on their scope and the limited new information or insights they contain, they are usually not appropriate for the journal. These long catalogues should certainly be available but in most cases they should be published, or — otherwise made accessible, by an appropriate government agency. It is refreshing to consider different types of manuscripts that will add a new dimension to the journal. I am pleased to report that we expect to initiate fairly soon a new series on the vascular plant flora of Canada. Information on this venture and the format for the accounts will be elaborated on by Associate Editor George La Roi in an upcoming issue of the journal. 1977 SMITH: EDITORIAL POLICY 119 Style Our suggested guide to authors regarding style is the CBE Style Manual. Webster’s Dictionary is our authority for English spelling. We request that all units of measure be expressed in the metric system and that authors use SI symbols. Other matters regarding The Canadian Field-Naturalist’s format and style can be ascertained by consulting the Instructions to Contributors or by looking over current journal issues. One subject about which the journal has never had a firm policy is whether the names of the authors of scientific names should accompany the scientific names in all papers. Our Editorial Board could not agree on this although the majority felt that the names of authors of scientific names should be included only for taxonomic or other papers where nomenclatural problems are involved. Therefore, unless an author is adamant, we intend to delete references to authors of scientific names except for taxonomic or other papers where the names are ina state of flux, as they are for some plant groups, many invertebrates, and many microorganisms. If a paper treats many species, a reference for the scientific names should be cited. We prefer authors to use “proper” common names for plants and animals throughout their papers unless there is a specific reason not to do so. Normally we have not capitalized the initial letters of common names, except in the case of birds where there is a list of capitalized “official” or “proper” names accepted by a learned society, the American Ornithologists’ Union; only certain insect names are capitalized in accordance with the list approved by the Entomological Society of America. We suggest that the initial letters of common names of other species of organisms should also be capitalized but at the moment will leave the decision on this up to individual authors. A species should be identified the first time it is mentioned in the abstract and in the text, preferably by putting the scientific name in parentheses following the common name. Generic names are written out in full the first time they are mentioned and thereafter may be abbreviated if it is unequivocally clear to which genus the initial refers. For consistency in nomenclature in non-taxonomic papers, we have adopted a policy for authors to follow certain mandatory or recommended references for the scientific and common names of organisms. For names of mammals and birds we require authors to use the following references. Mammals Jones, J. Knox, Jr., Dilford C. Carter, and Hugh H. Genoways. 1975. Revised checklist of North American mammals north of Mexico. Occasional Papers, The Museum, Texas Tech University 28: 1-14. Birds American Ornithologists’ Union. 1957. Check-list of North American birds. 5th edition, Baltimore. 691 pp. American Ornithologists’ Union. 1973. Thirty-second supplement to the American Ornithologists’ Union check-list of North American birds. Auk 90: 411-419. Also Corrections and additions to that supplement, 1973, Auk 90: 887. American Ornithologists’ Union. 1976. Thirty-third supplement to the American Ornithologists’ Union check-list of North American birds. Auk 93: 875-879. Also future supplements. Although the scientific and common names of other animals and plants seem to be less standardized than those of mammals and birds, the following list of publications may be helpful to authors as references for nomenclature. Amphibians and Reptiles Conant, Roger. 1975. A field guide to reptiles and amphibians of eastern and central North America. 2nd edition. Houghton Mifflin, Boston. 429 pp. Stebbins, Robert C. 1966. A field guide to western reptiles and amphibians. Houghton Mifflin, Boston. 279 pp. Fishes American Fisheries Society. 1970. A list of common and scientific names of fishes from the United States and Canada. 3rd edition. American Fisheries Society Special Publication 6. 150 pp. Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin 184. 966 pp. 120 THE CANADIAN FIELD-NATURALIST Vol. 91 Leim, A. H. and W. B. Scott. 1966. Fishes of the Atlantic coast of Canada. Fisheries Research Board of Canada Bulletin 155. 485 pp. McPhail, J. D. and C. C. Lindsey. 1970. Freshwater fishes of northwestern Canada and Alaska. Fisheries Research Board of Canada Bulletin 173. 381 pp. Hart, J. L. 1973. Pacific fishes of Canada. Fisheries Research Board of Canada Bulletin 180. 740 pp. Vascular Plants Hosie, R. C. 1969. Native trees of Canada. 7th edition. Queen’s Printer, Ottawa. 380 pp. Fernald, M. L. 1950. Gray’s manual of botany. 8th edition. American Book Co., New York. 1632 pp. Gleason, H. A. 1952. The new Britton and Brown illustrated flora of the northeastern United States and adjacent Canada. 3 volumes. (5th printing, 1974.) Published for the New York Botanical Garden by Hafner Press, New York. Volume 1, /xxv + 482 pp.; Volume 2, iv + 655 pp.; and Volume 3, iii + 595 pp. Manuscripts Before submitting a manuscript to The Canadian Field- Naturalist, authors should be certain that the publication is not premature, this journal is the proper outlet for their work, the contribution is a significant and new contribution to the scientific literature, the biology is emphasized, and they have followed the Instructions to Contributors. Their study, as reported, should be reasonably complete and comprehensive. Often manuscripts would benefit from more peer review before submission. The journal tries to restrict the papers published to natural history ones of national interest, 1.e., papers with primary (new) information or with new insights. Therefore, authors whose papers are mainly of local interest are advised to submit their papers to regional journals. Certain specialized subjects, e.g., wildlife management, statistical studies, pollution monitoring, detailed anatomical studies, extensive lists of the flora and fauna of an area, or papers that rehash earlier ones are also best submitted elsewhere. It is rare for a manuscript to be accepted for publication as it is submitted. Sometimes a paper, returned to an author for revision, is resubmitted without the requested changes and with no explanation or rationale for ignoring the criticisms. It is then impossible for an editor to judge sucha paper fairly. Other times an author will state the importance of his research in a letter. One can only ask the author, “Why didn’t you say that in your manuscript?” Most submitted manuscripts are too long. With the current expansion of the scientific literature, journals should publish only clear and concise write-ups; all unnecessary data, detail, and discussion must be deleted. Material that is supplementary to the published text may be directed to the Depository of Unpublished Data. A few suggestions and comments are offered to authors here because the same faults and shortcomings recur so often. Make the paper’s title descriptive and the sequence of the paper logical and orderly. Include a clear statement of purpose, rationale, and scope of the study in the Introduction. Put the study in the proper perspective with what is already known on the subject but limit the literature review to defining the problem at hand. Avoid extreme speculation with no basis in fact, subjective observations, excessive use of “weasel” words, and unwarranted conclusions. When citing a personal communication give the person’s initials and address. In the Results and Discussion make sure your statements are supported by data, the number of significant figures in the numerical data is not greater than warranted by the precision of your experimental method, and the work reported is reasonably complete and comprehensive. Point up the significance of your own results and limit the space devoted to outlining the results of others. Be careful with your terminology. For example, although “parameter” is often used indiscriminantly to mean any variable that can be measured, its original meaning restricts the term to a quantity that in any particular case is fixed or constant. Thus in the simple equation y = a + bx the parameters are a and b while the variables are x and y. The number of figures and tables should not be larger than is necessary to convey the message. Figure captions and table legends should give the purpose of the figure or table, and not just point out what they contain. Many figures require a scale, e.g., a scale line or bar to indicate a distance. 1977 SMITH: EDITORIAL POLICY 121 This reference line should be placed directly on the figure itself, not in the caption, so that it will be valid when the figure is reduced for printing. Graphic material should stand on its own without reference to the text. If original figures are larger than a normal page, it would make life easier for us if they were submitted as page-size (or smaller) photographic reproductions. Some authors write their acknowledgments with everyone in mind. It is logical to acknowledge the sources of financial aid, data, and illustrations, and to thank those who have given considerable help with the research or writing. Others, however, who have helped with the paper ina broad sense, or in a minor way, should remain anonymous. Normally, as titles before people’s names do not add anything, we delete them. Invariably some of the references in the Literature Cited don’t agree with those in the text and often some are incomplete. Careful proofreading beforehand would avert these errors. Unfortunately we do not have the staff and facilities available, as do some journals, to check out all the references. Because of this and because of the number of disciplines we cover, we have found it easier to ask authors to write out the names of journals in full despite the common use of abbreviations elsewhere. Nevertheless, many authors would prefer to use standard abbreviations. Therefore, it seems wise to give this a trial. Consequently we will allow authors to abbreviate names of serials and request that they follow the abbreviations listed in the Bibliographic guide for Editors and Authors. The onus must be on the author to ensure that the abbreviations used (as well as dates, pages, etc.) are correct because in many cases we will be left with little choice but to publish them as submitted. If there is any doubt about clarity, then the references should be written out in full. Usually initials are substituted for the given names of authors in the Literature Cited. Nevertheless if authors wish it, we are willing to publish the given names provided they were used originally. With the ever-increasing variety of methods of replicating and distributing written works, it is becoming increasingly difficult to know if, or how, to cite them. The members of our Editorial Board have agreed that we should retain Literature Cited and not change to References Cited or References. It was felt that the essence of good science is that anyone should be able to check not only the conclusions from a quality paper but also the references. Literature Cited comprises works that have been referred to in the text and that have been published, scheduled for publication (please note that this does not apply to manuscripts that have been merely submitted to a journal but not yet accepted), or deposited in libraries as theses; i.e., they are readily available. It does not contain unpublished references such as reports, memoranda, data sheets, internal documents, projects by university honor students, agency reports, consultants’ reports etc. Some of these exist only as a few copies while others are reproduced for limited distribution as xerographic copies or mimeographed sheets but in general most are not readily accessible. Even if the report is more widely distributed, e.g., nest record scheme reports, we discourage their use. In any case they are not literature and hence can be referred to only in the text. Page references in a book can be of value when they are given in the text for specific points. But how useful is it to conclude in a reference the number of pages in a book? In the past we have asked authors to include the total pagination. Although this may be useful as a guide to finding a book on library shelves or perhaps helps to give a reader an idea about the scope of the book, the number of pages does not seem otherwise of value; henceforth we will not specifically ask authors for this. We will, however, include this information if it is provided. I conclude this editorial with the thought that we have an open mind regarding many editorial policies. If we are convinced that it is time to change, revamp, or otherwise alter our policies, then we will do so. Responses of our authors and readers to this editorial or to any other matters concerning the publication of The Canadian Field-Naturalist are welcomed. LORRAINE C. SMITH, Editor Distribution and Abundance of the Prairie Rattlesnake, Crotalus viridis viridis, in Canada GEORGE B. PENDLEBURY 304 Manora Rd., N.E., Calgary, Alberta T2A 4R6 Pendlebury, George, B. 1977. Canadian Field-Naturalist 91(2): 122-129. Distribution and abundance of the Prairie Rattlesnake, Crotalus viridis viridis, in Canada. Abstract. The Prairie Rattlesnake, Crotalus viridis viridis, reaches the northern limit of its range in southeastern Alberta and southwestern Saskatchewan. The dominant limiting factor is likely the presence of a dry steppe climate. But, within this climatic zone, bedrock geology has played an important role in localizing den sites, and indirectly, in providing a suitable substrate. With greater human population pressure, and particularly the increased use of irrigation, the range of C. v. viridis has decreased over the past few decades. Prairie Rattlesnakes are most abundant in, and in close proximity to, major river valleys where irrigation is usually not practised, and where suitable den sites are developed. ‘The widely distributed Prairie Rattlesnake, Crotalus viridis viridis, reaches the northern limit of its range in southern Alberta and Saskatchewan. Klauber (1956), Wright and Wright (1957), Conant (1958), and Stebbins (1966) included portions of these provinces in their range maps for the species. Williams (1946), Logier and Toner (1961), Lewin (1963), Cook (1965), and others have reported Cana- dian capture locations. This study was undertaken in an attempt to define better the range of C. v. viridis in Canada, and reveal some of the factors which influence its distribution. The sources of locality data used were (1) museum specimens, (2) published reports, (3) my own observations and col- lections, (4) answers to mailed questionnaires, and (5) reliable reports made to others. Method Rattlesnakes are distinctive (Figure 1) in many respects and not generally confused with the other snake species which occur in Alberta and Saskatchewan. For this reason, I believed it might be possible to supplement existing distribution data by mailing a questionnaire to towns within the study area. The questionnaire pertained only to rattle- snakes and was organized so that most of the questions could be answered by marking an ‘X’ in the appropriate space after each question. The questionnaires and stamped, self-addressed envelopes were mailed to the postmasters of just over 100 selected towns in southern Alberta and Saskatchewan. Ninety-nine replies were received. Several advantages of this approach were that (1) the questionnaires could be mailed to a specific person in each town, (2) the organiza- tion of the questionnaire and the stamped, self- addressed envelope required no expense and little time on the part of the recipient, (3) the questionnaires were received, answered, and promptly returned by the postmasters in their respective post offices, and (4) each reply was likely based on the collective knowledge and experience of several individuals since rural postmasters generally come in contact with most of the local people. The main disadvantages inherent in data acquired with this method were (1) the imprecise locality data, and (2) the subjectiveness of replies concerned with abundance. In spite of these drawbacks, the data showed an amazing degree of coherence when plotted and analyzed. Distribution of Prairie Rattlesnakes Climatic changes subsequent to the end of Wisconsin glaciation 11 000 years ago fostered the establishment of prairie grassland in the southern parts of Alberta and Saskatchewan, continuous with the Great Plains of the United States. The southern grassland fauna moved northward into Canada along river valleys such as the Missouri and its tributaries. Once in Alberta, dispersion occurred along the valleys of existing rivers as well as those of abandoned meltwater channels (Williams 1946). Routes 122 1977 PENDLEBURY: THE PRAIRIE RATTLESNAKE IN CANADA 123 FIGURE |. The typical striking coil of a prairie rattlesnake. As part of a long-term project, color-coded vinyl discs have been affixed to the second proximal segment of the rattle to facilitate positive identification of this specimen froma distance. Photographed at a den site 4.6 km west of Redcliffe, Alberta. likely followed by rattlesnakes are shown in Figure 2. Naturally occurring populations of C. v. viridis in Canada are restricted to the prairie and foothills grassland vegetation zones of Alberta and Saskatchewan. These zones encompass, with the exception of upland areas, most of the southern regions of these provinces. The climate of this area is classified as moist to dry steppe (Longley 1972). The limit of continuous distribution of the Prairie Rattlesnake in Canada is shown in Figure 3. In Saskatchewan the range consists of two components separated by 150 km. The southern portion includes the area south of the Cypress Hills, Wood Mountain highlands which extend from the Alberta-Saskatchewan border to just west of Killdeer. The northern segment follows the South Saskatchewan River a distance of 60 km from the Alberta—Saskatche- wan border to a point south of Eatonia. The range of the Prairie Rattlesnake in Alberta is continuous and joins the two Saskatchewan segments. The limit extends west to a longitude of 112°53’00” and north to a latitude of 51°22’00”. Two particularly in- teresting aspects of the range in Alberta are that the lobes extend 55-65 km to the northwest and west from the main part of the range, and that three of these lobes are bisected by major rivers. 124 Sees LES THE CANADIAN FIELD-NATURALIST Vol. 91 UDM2YD}DYSOS Saskatoon FIGURE 2. Dispersal routes likely used by Crotalus v. viridis. Shading represents areas with an elevation in excess of 1000 m; stippling represents the area of dry steppe climate (after R. W. Longley 1972). From south to north these are the Oldman, Bow, and Red Deer Rivers. Possible reasons for this curious distribution will be discussed later. Records from outside the Present-day Continuous Range A number of locality records outside the present-day continuous range are indicated in Figure 3. The most northerly occurrence in Alberta is based on the 1943 report of a rattlesnake found on a farm near Trochu. The snake was apparently under a window and rattled as a cowhand was going to lean over the sill. The animal’s life was spared but no other rattle- snakes have been reported from this area(M. J. Hampson, personal communication). The prox- imity of this occurrence to Ghostpine Creek, a tributary of the Red Deer River, and a Drumheller occurrence lead me to believe this is a valid record, and not a result of an accidental introduction. The Drumheller record is based on a specimen killed in July 1957 at a point 0.8 km upstream from the bridge over the Red Deer River at Drumheller (C. F. Everts, personal communication). The locality 100 km east-southeast of Calgary is based on Fowler’s (1934) statement that the Prairie Rattlesnake is found 80.5 km east of High River. An interesting newspaper account from 1934(?) mentions that a young rattlesnake was found basking on a sandbar at the junction of Sheep Creek and the Highwood River. This location is 21.6 km south-southeast of Calgary. R. L. Fowler states in a letter to C. L. Patch dated 9 October 1934 that he was unable to verify the account but took it to be true. Rattlesnakes have been found, although rarely, in the vicinity of Saskatchewan River Crossing, south of Matador. Cook (1965) suggested that these occurrences might be the result of downstream rafting on debris. The records from Cypress Hills, Belanger Creek, and Eastend, reported by Logier and 1977 ie a a PENDLEBURY: THE PRAIRIE RATTLESNAKE IN CANADA 125 UDM2YIJD9SDS Saskatoon Sat a a Boe Pare ae OR ee USA FiGuRE 3. Data used to determine the distribution of Crotalus v. viridis in Canada. The solid line represents the present-day limit of continuous range; the dashed line is the limit according to Stebbins (1966). Symbols are as follows: solid circles = museum specimens and literature reports; half-solid circles = personal observations and collections; circle with dot = occurrences based on reports to sources othersghan myself; circle with cross = positive responses to the questionnaire and other occurrences reported to the writer; open circle = negative answers to the questionnaire. Toner (1955, 1961), were based on information obtained by L. M. Klauber during 1935-1938 by means of a postcard questionnaire. In reply to my questionnaire, the postmasters of Cypress Hills and Eastend indicated that rattlesnakes are absent in their respective areas. Hence, either the range has decreased or these occurrences were the result of chance migrations up the valley of the Frenchman River, a tributary of the Missouri River. All records from outside the present-day continuous range are at least 19 years old, and localities furthest from the limit are amongst the oldest. It is unlikely that these occurrences represent migrations from den sites. The meagre evidence available from Alberta indicates that seasonal movements are generally less than 4.3 km. Therefore, it is likely that the range of C. v. viridis in Canada is shrinking. Dinosaur Provincial Park at Steveville, Alberta, boasts of not having any rattlesnakes, yet the species was abundant here in 1912 (G. Lancaster, personal communication). Accidental translocation of C. v. viridis is not uncommon and a number of examples were brought to my attention during this investiga- tion. Mrs. O.M. Schafer of East Coulee, Alberta, wrote to me in 1972 “A live rattlesnake came in on a boxcar from eastern Alberta about 15 years ago.” The University of Alberta has a specimen that is suspected of having arrived at Edmonton in a load of hay from Montana (National Museum of Natural Sciences records). Another rattlesnake was found at a petroleum exploration drill-site just south of Edmonton. This animal had apparently crawled into some pipe lying on the ground when the rig was previously drilling in the Medicine Hat area (W.E. McKay, personal communication). A correspondent from Shaunavon, Saskatchewan, 126 wrote that “some have been known to come in with shipments of Alberta hay.” None of the accidental movements are plotted in Figure 3. Factors Influencing Distribution The distribution of C. v. viridis in Canada corresponds very closely to the dry steppe climatic zone (Figure 2) and it may be that climate is the dominant limiting factor. But I believe that, at least locally, there are other important controls. In my opinion, the bedrock and surficial geology has always exerted con- siderable influence on the distribution of C. v. viridis in Canada while, in the last two decades, a changing pattern of land use has contributed to significant changes. The curious lobed pattern of the range in Alberta may be due simply to the fact that the valleys of rivers bisecting these lobes were used as dispersal routes but bedrock geology is also involved. Figures 3 and 4 demonstrate the great degree of similarity in areal distribution of the Prairie Rattlesnake and bedrock of the Upper Cretaceous Belly River Formation. Asa result of Dyaqiy . : & ' = ? w ‘ es ' be ey } : Bb ok " ‘ x ee. * " < : : ’ ‘ NN J aN ., = .. Si 1 \ 1 ‘ ee ee a an rc mn nD a Co am sem sem msm mcm ems xen em AS is wm mn Se ED SS USA THE CANADIAN FIELD-NATURALIST UDM949}D¥SDS Vol. 91 fluvial erosion into nearly horizontal strata, the Belly River outcrop pattern tends to follow valleys as evidenced by the sections along the Milk, Oldman, Bow, and Red Deer Rivers. The bedrock pattern also emphasizes a number of abandoned channels. The rocks which constitute these strata are dominantly siltstones and very fine-grained sandstones with subordinate shales. The porous and poorly indurated sandstones absorb surface water readily and the interbedded shales act as gliding planes when saturated (Williams and Dyer 1930). Slumping is common along coulees, stream courses, and other places of pronounced relief (Powers 1931). The resulting fissures and caverns provide sites suitable for hibernacula, particularly where modified and enlarged through the burrowing activities of small mammals. Belly River sandstone outcrops contributed a large amount of material to overlying glacial deposits (Williams and Dyer 1930). The result- ing well drained sandy subsoil forms a ‘dry belt’ which “.. . is nearly coincident with the outcrop of the Belly River Formation, but extends Saskatoon Bedrock of Foremost and Oldman Formations Eel Irrigated Areas 105 FIGURE 4. Areas of Belly River (Foremost and Oldman) bedrock, and areas under irrigation in Alberta. 1977 farther south where glaciation has distributed the Belly River sands over the Pierre Bearpaw shale” (Williams and Dyer 1930). This soil likely forms a more favorable substrate for C. v. viridis. The surrounding bedrock is Bearpaw shale which has a tendency to“. . . weather to banks of earthy appearance” (Williams and Dyer 1930). Except where covered by Belly River sands, the soils overlying the Bearpaw are relatively poorly drained and thus provide a less suitable substrate. The hiatus separating the two Saskatchewan segments of the range was thought by Cook (1965) to result from a lack of available den sites. This may very well be the case, and may be related to bedrock geology. Most of the area is underlain by Bearpaw shale. Although the Belly River is present around the southern and eastern margins of a large Bearpaw outlier that straddles the Alberta-Saskatchewan border (Figure 4) the subdued topography results in few bedrock exposures and an absence of slump structures which play a significant role in den localization. Another factor contributing to the hiatus is the presence of unsuitable habitat associated with the Cypress Hills- Wood Mountain trend. These physiographic features rise toa maximum of 730m above the general level of the surrounding prairie. These landforms affect local weather conditions to the extent that precipitation and water supply are more abun- dant in their vicinity (Williams and Dyer 1930). These local variations in climate are naturally accompanied by changes in vegetation. Only eight occurrences within the present-day con- tinuous range were at elevations in excess of 1000 m (Figures 2 and 3). Changing land use is another factor which is exerting an important effect on rattlesnake distribution, particularly in Alberta. Increasing amounts of rangeland are being turned over to cultivation, and irrigation is required in many areas. Increased agricultural activities caused the Prairie Rattlesnake to be pushed out of parts of Kansas and Nebraska, and irrigation of California’s Imperial and Coachella valleys resulted in decreased numbers of Sidewinders, Crotalus cerastes laterorepens (Klauber 1956). It appears that C. v. viridis is suffering the same fate in Alberta. PENDLEBURY: THE PRAIRIE RATTLESNAKE IN CANADA 127 Comments from two of my correspondents are noteworthy. “Since the irrigation arrived in this area, this has gradually pushed the snakes into the coulees along the South Saskatchewan River and south into the Forty Mile Coulee” (A. K. Bateman, personal communication). “It is common belief rattlesnakes do not live in irrigation districts . . . Princess is lease land (dry pasture) and there are a few more seen in this area than Patricia” (Postmaster at Princess, personal communication). Imagery acquired by the earth-orbiting ERTS-I satellite was used to map areas under irrigation (Figure 4). Synoptic coverage com- bined with adequate resolution conferred an advantage on imagery over conventional aerial photography. The signature of infrared radia- tion reflected from vegetation is strongly dependent on plant type and vigor. Hence, the 0.7—0.8 uw (near infrared) band was found to be particularly useful for differentiating between irrigated areas under intensive cultivation, and areas of grain crops and rangeland. Figures 3, 4, and 5 indicate that rattlesnakes are not always excluded from irrigated areas. This is particularly evident in the southwestern lobe of the range near Lethbridge. This apparent anomaly may be reconciled, however, if the area’s topography and bedrock exposure are considered. Coulees are numerous and deeply eroded, exposing Belly River strata. The intervening areas are irrigated and cultivated, but the coulees themselves are not. Thus, peninsulas of favorable habitat extend from the valley of the Oldman River into less hospitable environs. Similar situations exist elsewhere in the province. Abundance Questionnaire recipients were asked to indi- cate whether rattlesnakes, if present in the immediate vicinity of their respective towns, were rare, common, or abundant. As already noted, replies to such a query are necessarily very subjective, and yield results which are semi- quantitative at best. This is particularly true when they are not based on an organized census. There is, unfortunately, no way of assigning a meaningful numerical range of values to each of the categories. The results, when plotted and 128 Q ae = ¢« Sees temee en Seem THE CANADIAN FIELD-NATURALIST Vol. 91 UDM9YD}DSDS Saskatoon RELATIVE ABUNDANCE Rare Common Abundant 105 FIGURE 5. Abundance of Crotalus v. viridis in Canada. contoured, generated the pattern shown in Figure 5. The ‘abundant’ category straddles valleys associated with major rivers. Relative abundance decreases away from the river valleys and in an upstream direction through the distribution lobes in Alberta. An anomalous situation exists near the range limit in the southwestern Alberta lobe where rattlesnakes are said to be common. A possible explanation was discussed in the preceeding paragraph. The closest area of ‘abundant’ occurrence lies 35 km to the east. The intervening area of ‘rare’ occurrence results from the fact that tributary coulees are less common, and those present are not as deeply eroded, i.e., favorable strata are not exposed. Furthermore, this area lies within an irrigation belt (Figure 4). Fluctuations in numbers were mentioned by four correspondents. In Alberta, the postmaster at Ralston said, “This year [1972] rattlesnakes are rather scarce... guess it just has not been hot enough.” The postmaster at Wardlaw, however, believed rattlesnakes “ ... are becoming more common.” Contradictory opinions were also obtained from two points in Saskatchewan. The postmaster at Burstall wrote, “More than average were seen here in 1973,” while“...inthe vicinity where McEachern Post Office was, rattlesnakes are rare, they were more abundant a few years back” (local resident, personal communication). Conclusions The results of this investigation lead to a number of conclusions: 1. Although distribution of C. v. viridis in Canada is controlled by many factors, available evidence indicates that, after cli- mate, bedrock geology and land use are two of the more important controls. 2. Bedrock geology has contributed to suitable habitat in two ways: (1) the mechanical properties of the Belly River Formation lead to slumping with the concomitant formation of suitable den sites, and (2) soils derived from the Belly River sandstones are well OWT drained and form a more suitable substrate. 3. A change in land utilization from rangeland to irrigated cultivation and the attendant habitat modification is restricting C. v. viridis, more and more, to the immediate vicinity of coulees. 4. Increased pressure from the human popula- tion in the future will result in a continuous decrease in the range of the Prairie Rattle- snake in Canada. Acknowledgments I am indebted to F.R. Cook, National Museum of Natural Sciences, and M. J. Hamp- son, then of the Provincial Museum and Archives of Alberta, for providing distribution data. ERTS-1 imagery was kindly provided by R. G. Agarwal. Special thanks are extended to the 99 postmasters who answered and returned my questionnaire. W. B. Preston and L. Mol- insky read the manuscript and offered sug- gestions for its improvement. Literature Cited Conant, R. 1958. A field guide to reptiles and amphibians of Eastern North America. Houghton Mifflin Company, Boston, Massachusetts. 366 pp. Cook, F. R. 1965. Additions to the known range of some amphibians and reptiles in Saskatchewan. Canadian Field-Naturalist 79(2): 112-120. Fowler, R. L. 1934. Some amphibians and reptiles of the PENDLEBURY: THE PRAIRIE RATTLESNAKE IN CANADA 129 district around High River, Alberta, 1933. Canadian Field-Naturalist 48(9): 139-140. Klauber, L. M. 1956. Rattlesnakes: their habits, life his- tories, and influence on mankind. University of Cali- fornia Press, Berkeley and Los Angeles. 1533 pp. Lewin, V. 1963. The herpetofauna of southeastern Alberta. Canadian Field-Naturalist 77(4): 203-214. Logier, E. B.S. and G. C. Toner. 1955. Check-list of the amphibians and reptiles of Canada and Alaska. Con- tributions of the Royal Ontario Museum of Zoology and Palaeontology 41: 1-88. Logier, E. B.S. and G. C. Toner. 1961. Check-list of the amphibians and 0.05; f-test); however, summer (11 June—9 September) flight initiation was significantly later than in either the spring (P< 0.01; f-test) or autumn (P< 0.05; t-test). Flights tended to begin earlier in relation to sunset on overcast compared to clear evenings (P>0.05; t-test) (Table 1). Duration of Activity During the breeding season, when displaying birds were in the air, it was difficult to determine the duration of evening flights from woods to fields. In the post-breeding period this was more easily done. On 46 evenings, when more than one woodcock was observed in the air, the period of flight activity for the population lasted 1-15 min and averaged 6.1 + 3.5 min (Table 1). Length of flight activity did not differ significantly (P> 0.05; t-test) over the season or under varying cloud conditions, although flights were of slightly longer duration on clear evenings. TABLE 1—Initiation and duration of woodcock flight activity in relation to cloud cover and season Flight initiation Duration of evening after sunset flight! (minutes) (minutes) Dates Overcast? Clear} Overcast Clear 21 April-10 June 17.8 + 11.0 25.9 = 11.3 (N =8)* (N = 13) 11 June—9 September 31.6 + 12.7 36.7 + 6.7 5,2) ae 3.3) 6.4+ 4.1 (N = 20) (N = 21) (N = 9) (N = 14) 10 September—28 October 22.0 + 7.8 29.1 + 5.4 S2ee 23 8.0 + 4.3 (N = 15) (N = 13) (N = 10) (N = 8) 'When more than one woodcock was observed per evening. 20.75 to full cloud cover. 30.0 to 0.25 cloud cover. 4Number of evening flight periods in sample. 1977 2030 2010 1950 1930 = Y 1910 LJ oi 1850 = - 1830 1810 & SUNSET 1750 1730 1710 Alle || LOM ZOWS OFS) APRIL MAY JUNE WISHART AND BIDER: WOODCOCK ACTIVITY, SOUTHWESTERN QUEBEC 3/4 TO FULL CLOUD COVER 6 1/4 TO 3/4 CLOUD COVER O TO 1/4 CLOUD COVER SAS) eS) JULY 143 IS) 28s 28 CMe. 2h. 6 62,28) Ss Ms AUG. SER OCT. NOV. FiGuRE 1. Seasonal relationships of woodcock flight initiation to sunset and cloud cover. Between 20 April and 4 June male woodcock performed courtship displays each evening, characterized by vocalizations (peents) on the ground, interspersed with aerial displays over the territory (Mendall and Aldous 1943). From first to last peent each evening, courtship lasted for 46.2+ 13.0 min (13 evenings). Sporadic peenting continued, however, on bright moonlit nights. Seasonal Changes in Flight Activity After 10 June, nightly counts were made of the number of woodcock seen and heard flying to openings from diurnal coverts. To avoid duplication, those performing protracted court- ship were not included. This index of activity showed that a distinct seasonal pattern occurred. Groups of values (Figure 2) were compared using the Mann-Whitney U test to detect the significance of changes. The summer peak in flight activity occurred between 27 June and 19 July. A significant decrease (P< 0.05) followed until 13 Septem- ber, which was in turn followed by an increase (P <0.01) that lasted until 6 October. Activity decreased thereafter and declined to zero by 29 October. 144 200 180 160 140 120 100 80 60 °° CUMULATIVE NUMBER OF WOODCOCK 40 . 20 OF UNE 27 a <_ onal JUNE 26 THE CANADIAN FIELD-NATURALIST Vol. 91 CONG Ip. As) OCT. 28 a OC Te sSiteater ees . OCT: 6 TOK lO, 2% 4 16 26" 8 14 JUNE JULY I) 25 2 AUGUST is 2 42 jO 1) 1 5 SERIT: OG IE. NOV. FIGURE 2. Cumulative frequency curve showing the number of woodcock flying to nocturnal fields each evening over the season. Arrows indicate the dates at which slopes change. The incidence of protracted courtship and mist-netting success similarly indicated a drop in woodcock activity in August followed by an increase before autumn migration. In June and July, courtship was heard on 21% of observed evenings (24 evenings) and in September and October this rose to 39% (34 evenings). In August, however, the incidence of protracted courtship was low (6% of 16 evenings). Mist- netting success also declined to a low of 0.4 captures per mist-net night during August (Figure 3). Diurnal habitat surveys were used as an index of woodcock density in the area and provided a means of determining what factors affected changes in flight activity. Between the spring and summer, surveys showed the woodcock popu- lation changed little (P > 0.05; chi-square test) in size. But after mid-September, a significant (P< 0.01; chi-square test) increase occurred. After 1 November, few woodcock remained in the area and none was flushed from previously occupied coverts in 174 min of survey (Table 2). Nocturnal Activity in Fields The sand transect, read over 26 days, yielded 1977 CAPTURES / MIST NET NIGHT APR. MAY JUNE WISHART AND BIDER: WOODCOCK ACTIVITY, SOUTHWESTERN QUEBEC 145 JULY AUG. SEPT. OCT. FIGURE 3. Number of woodcock captured per mist-net night over the season. Numbers in brackets indicate the number of mist-net nights recorded per month. High values in April and May are due to the site tenacity of territorial males. 86 woodcock crossings. Distinct crepuscular peaks of activity occurred within 1h of the estimated periods that woodcock entered and departed from fields at that time of year. No crossings were recorded before 1800, between 2100 and 0400, or after 0600 hours Eastern Standard Time. Evening activity (76 crossings) was 7.6 times greater than that in the morning (10 crossings). TABLE 2—Woodcock flush and sign surveys in diurnal habitat indicating seasonal changes in density Woodcock Woodcock Hours of flushed sign per Dates survey per hour hour 20 April- 10 June 36.9 0.7 8.9 11 June- 9 September 36.0 1.0 9.0 10 September- 1 November 15.8 2.0 28.5 Discussion The timing of evening flights from diurnal to nocturnal cover changed over the season. Therefore, it seems that flight initiation is governed by a critical light intensity (Duke 1966) which is altered over the year in its time of occurrence because of the change in rate of the sun’s descent. The duration of twilight varies with the constant change in position of the earth in relation to the sun (McGraw-Hill Almanac 1969). The slower descent of the sun in June and July, therefore, is likely the reason for the skewed nature of Figure |. Leopold (in Welty 1962) observed a similar pattern of commence- ment times of morning song by several species. Greater cloud cover tended to be correlated with earlier flight initiation, but the relationship was not significant (0.1 > P > 0.05; t-test). Thus, although woodcock did respond to seasonal changes in light conditions, day-to-day varia- tions caused by cloud were not enough to elicit a large response. In Maine, Krohn (1971) ob- served a similar relationship. 146 Evening courtship displays averaged about 46 min, which agrees with findings by Duke (1966) studying woodcock at a similar latitude. In Maryland, however, Sheldon (1967) found courtship lasted for only 20 min and concluded that birds in northern areas perform longer. Miller (1958), studying the American Robin (Turdus migratorius), reported similar findings concerning length of morning song. This relationship suggests that latitudinal differences of crepuscular light conditions are responsible for the observed variability. As one moves southward on a given day (in the spring) there is an increase in the rate of the sun’s descent at night and ascent in the morning (McGraw-Hill Almanac 1969). Thus, the length of time each day with light intensities typical during wood- cock courtship is shorter further south. It is not known if this variability of length of courtship affects woodcock productivity at various lati- tudes. All three indices conclusively indicated a decline in flight activity of woodcock in August, followed by a rise in autumn (see also Sheldon 1961; Krohn 1971). Despite these changes in flight activity and recruitment of juveniles, diurnal habitat surveys showed the population was almost stable in size over the spring and summer. Autumn surveys, however, indicated an increase in the population, likely as a result of more northern migrants moving into the area. There- fore, at least part of the rise in activity after mid- September can be attributed to the rise in woodcock density. Flight activity declined after 8 October and reached zero on 29 October, indicating that migration from the area was completed. Woodcock undergo a complete post-breeding molt and the nutritional demands of feather growth may be the reason for the mid-summer waning in activity. Heinroth and Heinroth (1958, p. 91) stated that when birds molt they are less active, energy being used to renew feathers. Payne (1972), reviewing the literature, indicated that metabolic rate of molting birds may increase up to 30% over that of non-molters. Owen and Krohn (1973) found the peak of the woodcock molt in Maine occurred in August. Fat deposition was negatively correlated with the molt and they judged that most of these birds were physiologically unprepared for migration THE CANADIAN FIELD-NATURALIST Vol. 91 until mid-October. Thus, the August decline in flight activity of woodcock is correlated with the peak of physiological stress due to molt. At this time a portion of the population seems to remain in forested habitat throughout the day and night. It is unknown if all individuals follow this pattern, at what stage of stress reduced activity occurs, and over what length of time it persists for an individual. Throughout the summer and autumn, birds of all sex and age classes were captured on fields at night, but data were insufficient to determine if a particular class was most responsible for the decline in activity. Woodcock not participating in crepuscular flights and remaining in coverts at night would conserve energy. Earthworm surveys have shown that relatively few food items are found in summer fields (Wishart and Bider 1976) and little feeding occurs there (Krohn 1970). During the molt, birds may tend to remain in coverts where food is plentiful, and actively feed throughout the day and night there until a level of physical condition is re-attained. Sand transect information showed that wood- cock were inactive for most of the night after they had entered fields. Peaks of activity occurred within one hour of entrance and departure. Dunford and Owen (1973) and Owen and Morgan (1975), using birds equipped with radio transmitters, also found them sedentary for most of the night. Sheldon (1961) believed that woodcock flew to fields to feed, as occurs on the wintering grounds (Glasgow 1958). Krohn (1970), however, found that little food was obtained on summer fields, and Dunford and Owen (1973) speculated that they may act as roosting sites, providing security from preda- tors. Unfortunately little is known about predation on woodcock, and the advantage to birds of roosting in open areas is not clear. Bider (1968) has shown that few potential woodcock predators consistently hunt in open fields. At the study area, activity on fields took place soon after birds entered and just before morning departure. Krohn (1971) and Wishart and Bider (1976) observed that woodcock dispersal on summer fields was not random. If fields are indeed used for roosting, birds may have to search for sites with suitable microhabitat cover before settling for the night. This may require them to make several flights over the area, to Sy change fields a number of times, and to do some searching on the ground. Why then are birds also active prior to morning departure? Mendall and Aldous (1943) believed early morning and late evening were favorite feeding times for wood- cock. Krohn (1970) pointed out that whereas little food was obtained on fields there was evidence that some foraging took place there. The morning surge in activity, therefore, may be due to foraging. The lack of success in finding sufficient food then might in part stimulate birds to return to diurnal habitat where both food and cover are more abundant. Acknowledgments We thank the assistants who read the sand transect in 1967 and 1968. W. B. Krohn, R. W. Stewart, W.L. Vickery, and J.G. Doucet offered many helpful suggestions, and G. S. Hochbaum and S. G. Sealy reviewed different drafts of the manuscript. This study was funded by a grant from the National Research Council of Canada. Literature Cited Bider, J. R. 1968. Animal activity in uncontrolled ter- restrial communities as determined by a sand transect technique. Ecological Monographs 38: 269-308. Duke, G. E. 1966. Reliability of census of singing male woodcock. Journal of Wildlife Management 30: 697-707. Dunford, R.D. and R.B. Owen, Jr. 1973. Summer behavior of immature radio-equipped woodcock in central Maine. Journal of Wildlife Management 37: 462-469. Glasgow, L. L. 1958. Contributions to the knowledge of the ecology of the American Woodcock, Philohela minor on the wintering range in Louisiana. Ph.D. thesis, Texas A. & M. University, College Station. 153 pp. Greeley, F. 1953. Sex and. age studies in fall shot WISHART AND BIDER: WOODCOCK ACTIVITY, SOUTHWESTERN QUEBEC 147 woodcock (Philohela minor) from southern Wisconsin. Journal of Wildlife Management 17: 29-32. Heinroth, O. and K. Heinroth. 1958. The birds. University of Michigan Press, Ann Arbor. 181 pp. Krohn, W. B. 1970. Woodcock feeding habits as related to summer field usage in central Maine. Journal of Wild- life Management 34: 769-775. Krohn, W. B. 1971. Some patterns of woodcock activities on Maine summer fields. Wilson Bulletin 83: 396-407. Martin, F. W. 1964. Woodcock age and sex determination from wings. Journal of Wildlife Management 28: 287-293. McGraw-Hill Almanac. 1969. Directory and almanac of Canada. McGraw-Hill Book Company, Toronto. 864 pp. Mendall, H. L. and C. M. Aldous. 1943. The ecology and management of the American Woodcock. Maine Co- operative Wildlife Research Unit, University of Maine, Orono. 201 pp. Miller, R. C. 1958. Morning and evening song of robins in different latitudes. Condor 60: 105-107. Nelson, L. K. 1955. A pheasant neck tag. Journal of Wildlife Management 19: 414-415. Owen, R.B., Jr. and W.B. Krohn. 1973. Molt patterns and weight changes in the American Woodcock. Wilson Bulletin 85: 31-41. Owen, R.B., Jr. and J.W. Morgan. 1975. Summer behavior of adult radio equipped woodcock in central Maine. Journal of Wildlife Management 39: 179-182. Payne, R.B. 1972. Mechanisms and control of molt. In Avian biology. Volume II. Edited by D.S. Farner and J. R. King. Academic Press, New York. pp. 103-155. Sheldon, W. G. 1961. Summer crepuscular flights of Amer- ican Woodcocks in central Massachusetts. Wilson Bulletin 73: 126-139. Sheldon, W. G. 1967. The book of the American Wood- cock. University of Massachusetts Press, Amherst. 227 pp. Welty, J.C. 1962. The life of birds. Company, Toronto. 546 pp. Wishart, R. A. and J. R. Bider. 1976. Habitat preferences of woodcock in southwestern Quebec. Journal of Wild- life Management 40: 523-531. W.B. Saunders Received 11 March 1976 Accepted 7 January 1977 Reproductive Success of Herring Gulls Nesting on Brothers Island, Lake Ontario, in 1973 STANLEY M. TEEPLE Canadian Wildlife Service, Ottawa, Ontario KIA 0H3 Teeple, Stanley M. 1977. Reproductive success of Herring Gulls nesting on Brothers Island, Lake Ontario, in 1973. Canadian Field-Naturalist 91(2): 148-157. Abstract. Breeding success and causes of breeding failure were assessed in 1973 ina Herring Gull colony of 34 pairs in eastern Lake Ontario. Breeding synchrony was normal, 13 pairs laid repeat clutches, but 77% of all eggs laid failed to hatch. The number of chicks fledged per pair averaged at least 0.06 but not more than 0.18, an exceptionally low result. Geometric mean concentrations of DDE and PCBs in 15 eggs that failed to hatch were 134 and 420 ppm dry weight. Concentrations of dieldrin, p.p'DDD, p,p’DDT, heptachlor epoxide, B-benzene hexachloride, hexachlorobenzene, and mercury were each less than 6 ppm. Arithmetic mean shell thickness of 13 of those 15 eggs was 0.339 mm, and mean thickness index of 11 of those 13 was 1.60; both are low values. Pathological examinations and analyses for organochlorine pesticides and PCBs in brains were conducted on 12 chicks that died. For 11 of the 12, no clear cause of death could be determined. A general association was established between high organochlorine levels and the low breeding success. Studies by Keith (1966) and by Ludwig and Tomoff (1966) showed low breeding success and high organochlorine (OC) residue levels in populations of Herring Gulls (Larus argentatus) in Lake Michigan in the mid-1960s. Hickey and Anderson (1968) demonstrated a relationship between egg-shell thinning and levels of DDE in the eggs of five Herring Gull populations in eastern North America and later documented shell thinning in a number of North American bird populations, among them Herring Gulls on the Great Lakes (Anderson and Hickey 1972). They found that shell thinning in the Great Lakes population began in the late 1940s or early 1950s, and that during the period 1960-1969 shells were 10% thinner than in eggs collected prior to 1947. Gilbertson (1974) and Gilbertson and Hale (1974) also have shown that low breeding success coupled with thin-shelled eggs having high OC levels prevailed in Herring Gulls in the lower Great Lakes in 1972 and 1973. This paper reports on the breeding success, shell thicknesses, and OC residue levels of non- hatching eggs and chicks that died in a small Herring Gull colony in eastern Lake Ontario in 1973, and provides further documentation of the low reproductive success of Herring Gulls nesting in Lake Ontario. Study Area and Methods “Brothers Island” (44° 12’ N, 76°39’ W) is the most westerly of the three Brothers Islands in eastern Lake Ontario, 13 km west of Kingston and about 1.3 km offshore, between Amherst Island and the mainland. Brothers Island is approximately 215 m long and varies from | to 10 m in width. Elevation was about | m above the lake level prevailing in the 1973 breeding season. The gulls nested on a flat area of lime- stone running most of the length of the island. Over its full length, trees, shrubs, and climbing vines were interspersed among relatively open areas bearing only ground vegetation. Tangles of vines, shrubs, and driftwood provided hiding places for gull chicks, as did crannies in the broken limestone. Except for three pairs of Mallards (Anas platyrhynchos) and a single pair each of Song Sparrows ( Melospiza melodia) and Red-winged Blackbirds (Agelaius phoeniceus), Herring Gulls were the only species nesting on the island. I first inspected the colony in 1973 from the air on 9 April. Starting on 26 April, I visited the island by boat every 2 to 3 days during April, May, and June (except for one 6-day interval in May) and then every 4 to 5 days until 18 July, with a final visit on 29 July. The visits, usually by two observers, were from 30 min to 2 hin length, and averaged about | h. I marked all gull nests by writing a nest number in indelible felt pen on a nearby rock, which was also marked with a spot of orange paint to help in relocating the nest number. All eggs were numbered with a non-toxic felt pen 148 1977 when first found, and in the sequence in which they were laid, if known. I banded each chick when I first found it, using standard United States Fish and Wildlife Service metal bands. On each visit, I recorded the numbers and condition of nests, eggs, and chicks. Eggs that were no longer being incubated, or were out of the nest and had been out on several successive visits, were collected, as were egg-shells found outside the nests and all dead chicks. In the laboratory, the contents of each egg were emptied into an acetone-rinsed glass jar and examined to determine, when possible, stage of development at death. Chicks were examined for injuries and pathological conditions, including endoparasites, bacterial infection, and malnutri- tion that could account for their deaths. Egg contents and dead chicks were frozen. All egg-shells were air-dried for several weeks, after which the thickness of each shell was measured to the nearest 0.001 mm at five equi- distant points around the girth, using a Starrett Model 1010 dial micrometer with ball anvils. Measurements included the shell membranes. I calculated thickness indices (Ratcliffe 1967) for those shells for which weights and dimensions were available. All egg contents and brains were analyzed individually by gas-liquid chromatography for OC pesticide and polychlorinated biphenyl (PCB) residues at the Ontario Research Founda- tion. Each of the egg contents was also analyzed for total mercury. Moisture and fat percentages were determined for each egg and brain. The PCB values given are based on a reference mix- ture of Aroclor 1254:1260 (1:1). Analytical methods for OC pesticides and PCBs were des- cribed by Reynolds and Cooper (1975), and for mercury by Fimreite and Reynolds (1973). Results Breeding Biology I believe the number of breeding pairs of Herring Gulls (pairs that built at least one nest and laid at least one egg) on Brothers Island in 1973 was 34; that number is used in computing the various statistics of breeding success. It was determined from the number of first clutches among the total 47 clutches initiated. Thirteen of those 47 were thought to be second clutches, TEEPLE: HERRING GULLS, LAKE ONTARIO 149 although it was not possible to know that with absolute certainty, since gull pairs were not marked. It has been established, however, that Herring Gull pairs usually renest in the im- mediate area of the previous nest (Paludan 1951) and usually initiate the second clutch I1 to 17 days after loss of the first (Goethe 1937; Paludan 1951). On these bases, and having regard to the fate of each earlier clutch and to the spatial locations and times of initiation of the 13 clutches, I concluded that they were second clutches. | The gulls were already occupying territories on 9 April. On 26 April there were 34 nests, of which five were incomplete. Four clutches had been completed (i.e., contained no additional eggs on subsequent visits) and | 1 others started. The median date of initiation of first clutches was 28 April (Figure 1), and 68% of those clutches were initiated during the 12-day period 22 April to 3 May. Dates of initiation of the 14 clutches begun prior to my first visit were estimated by back-dating from the date of clutch completion or hatching. One clutch was com- plete at the time of my first visit, but failed to hatch; I am certain only that it was begun prior to 23 April. The median date of initiation of clutches regarded as repeat layings was 19 May (Figure 1). Sixty nests were completed during the season, of which 46 contained at least one egg, while the remaining 14 were abandoned or destroyed prior to the start of egg-laying (Table 1). Many of the abandoned/ destroyed nests were built close to the water’s edge, and were subject to the effects of wave action. One of the 46 nests with eggs had both a first and second clutch laid in it, the first clutch having disappeared during egg-laying; 12 other nests among the 46 are believed to have been renestings resulting from failure of the original attempt. Twenty-one nests contained eggs that hatched and 25, including 15 of the 21 with eggs that hatched, are known to have contained eggs that were infertile or had dead embryos. The total number of eggs found in the colony was 124, comprising 44 completed clutches and three that were abandoned or destroyed prior to completion. Of the 124 eggs found, 28, or 23%, hatched. Of the remaining 96 that failed to 150 5 UO oO ‘4 5 ic o Ore O ~ = ee O (oe 2 ae >) Ze O 15 20 25 30 April THE CANADIAN FIELD-NATURALIST 5 Vol. 91 eens al First Clutches Second Clutches 10 IS ZO) NS) May 30 FiGuRE |. Dates of initiation of 33 first clutches and 13 second clutches by Herring Gulls on Brothers Island in 1973. hatch, 38% were infertile or failed because of embryonic mortality, 17% disappeared from the nest, 25% disappeared when the entire nest was destroyed, and 21% failed for various other reasons, including destruction by adult gulls and TABLE 1—Outcome of Herring Gull nests on Brothers Island in 1973 Total nests started 63 Not completed 3 Completed 60 Contained no eggs 14 Contained eggs 46! Clutch not completed — abandoned during egg- laying, when eggs disappeared or were out of nest 2 Clutch completed, incubated 44 Abandoned during incubation when eggs dis- appeared, were destroyed, or were out of nest 7 Destroyed during incubation, probably by wave action 9 Eggs failed to hatch 7 Hatched chicks 21 'One of these nests contained two successive clutches, the first clutch having disappeared during egg-laying. rolling out of the nest. Table 2 details the fate of eggs in the colony. Hatching and fledging success in completed clutches is summarized in Table 3. Sixteen eggs were collected from the colony. They represented either 12 or 13 different clutches, two of which were second clutches. Two of the eggs were fresh and 14 were rotten. No visible embryonic development had taken place in either fresh egg. Of the 14 rotten eggs, two had been rolled from the nest during early incubation, one had been incubated 16 to 19 days, and I1 had been incubated at least 27 days. Among those 12 eggs that had been incubated for a significant period, four showed no dis- cernable embryonic development. Development in the remaining eight was estimated to be | to 25% in three, 51 to 75% intwo, and 76 to 100% in three. In all, there were 28 clutches (20 first clutches and 8 second clutches) in which at least one of the eggs in the clutch remained intact through 27 days of incubation. The fate of the 80 eggs found in those clutches is summarized in Table 4. Among the 61 eggs that were incubated 27 or more days, 28 (46%) hatched and 33 (54%) failed to hatch owing to infertility or embryonic 1977 TEEPLE: HERRING GULLS, LAKE ONTARIO 151 TABLE 2—Fate of Herring Gull eggs on Brothers Island, 1973 In 34 first In 13 second In all 47 clutches clutches clutches During egg-laying Disappeared from nest 7 0 q Out of nest l 0 l During incubation Disappeared from nest 6 3 9 Disappeared when nest destroyed, probably by wave action 24 0 24 Destroyed by adult gulls 3 5 8 Out of nest 6 3 9 Shell damaged l 0 I Embedded in nest 0 l 1 Infertile, or rotten early embryo not discernable 13 10 23 Embryo died before pipping 7 3 10 Pipped and died 3 0 3 Hatched 21 7 28 Totals 92 32 124 mortality. Those 28 clutches represented 28 of the 34 gull pairs in the colony. The 28 chicks hatched represent 0.82 chicks per breeding pair. Thirteen of those were later found dead. Two chicks were known to have fledged (were seen flying) and four more were seen at 17 to 35 days of age, and may have fledged. Thus, 2-5% of eggs in the colony produced chicks that fledged. Of chicks that hatched, between 7 and 21% fledged. In terms of chicks fledged per breeding pair of adults, these numbers represent 0.06 chicks known to have fledged and a maximum of 0.18 that may have fledged. Chick Mortality and Chemical Residues in Brains Of the 13 chicks found dead 1n the colony, 12 were collected. Ages at death ranged from a few hours to about 7 weeks. One chick, about | day TABLE 3—Clutches completed by 34 Herring Gull pairs on Brothers Island, 1973 Number of Number of clutches Number of Number of clutches hatching chicks chicks chicks hatched fledged First clutches! Two-egg 9 3 3 1(?) Three-egg 22 12 18 2-5 Both two- and three-egg 31 15 21 2-6 Second clutches! Two-egg 7 3 3 0 Three-egg 6 3 4 0 Both two- and three-egg 13 6 i] 0 Both clutches! Two-egg 16 6 6 1(?) Three-egg 28 15 22 2-5 Both two- and three-egg 44 21 28 2-62 'At start of incubation. 2From six clutches. 152 THE CANADIAN FIELD-NATURALIST Vol. 91 TABLE 4—Fate of Herring Gull eggs found in 28 clutches in which at least one intact egg remained in the nest for 27 or more days of incubation, Brothers Island, 1973 In 20 first In 8 second In all 28 clutches clutches clutches During egg-laying Disappeared 5 l 6 During incubation Disappeared 3 l 4 Destroyed by adult gulls 3 l 4 Out of nest 3 0 3 Shell damaged l 0 ] Embedded in nest 0 | 1 Incubated full term but failed to hatch 23 10 33 Hatched 21 7 28 Totals 59 21 80 old and near death, was in convulsions. Parasites were absent, and none of the chicks showed skull fractures to indicate it had been pecked by adult gulls. Six showed evidence of disease (four had diarrhoea, one had a very yellow liver, and one showed prominent convolutions in the kidney tubules); three, including two of the diseased ones, exhibited signs of malnutrition. No signi- ficant pathogenic bacteria were isolated from any of 10 from which tissues were cultured. As the chicks gained weight brain residue levels of the various OC chemicals declined rapidly, presumably as a result of the dilution effect of the increase in body tissue. Seven of the 12 chicks for which brain analyses were done weighed 56 g or less, and may be regarded as newly-hatched chicks. Among those seven, the geometric mean and 95% confidence interval for DDE, in parts per million (ppm) or milligrams per kilogram dry weight were 147 and 102-213; for PCBs, 651 and 491-864. Mean brain residues and confidence intervals in the other five chicks weighing 140-724 g were 27.7 and 10.9-70.4 ppm DDE and 135 and 53.6-341 ppm PCBs. Low levels of dieldrin, p,p’ DDT, heptachlor epoxide, and hexachlorobenzene were also detected in each of the 12 brains. In the brains of the seven small chicks, concentrations of each of the above chemicals were less than 7 ppm dry weight and in the five larger chicks, less than 2 ppm. Residue levels on a wet weight basis in the seven smaller and five larger chicks may be calculated by multiplying the dry weight values by 0.14 and 0.16, respectively. Chemical Residues in Eggs and Shell Thickness The contents of each of 15 eggs were analyzed for OC pesticides, PCBs and total mercury. Geo- metric means and 95% confidence intervals for DDE and PCBs in those 15 eggs are compared in Table 5 to similar measurements from eggs collected in 1972 in Lakes Huron (Georgian Bay), Erie, and Ontario, and the Gulf of St. Lawrence and the Bay of Fundy, New Bruns- wick (Gilbertson and Reynolds 1974). The arith- metic mean and 95% confidence interval for DDE ona dry weight basis in those 15 eggs were 144 and 110 to 178, and for PCBs 456 and 345 to 567. Since the eggs analyzed in this study had partially dehydrated prior to collection, it was not possible to calculate a factor for converting residue levels from a dry weight to a wet weight basis. However, the mean conversion factor for 160 fresh Herring Gull eggs from the Great Lakes was 0.24 (standard deviation 0.01) (inter- nal Canadian Wildlife Service report from the Ontario Research Foundation). Approximate residue levels on a wet weight basis may be obtained by multiplying the dry weight levels by that factor. Dieldrin, p,p’DDD, p,p’DDT, heptachlor epoxide, B-benzene hexachloride, hexachloro- benzene, and mercury were also found in each egg, but in every case at concentrations less than 6 ppm dry weight. Thirteen of the eggs collected had shells that remained sufficiently intact to allow shell thick- ness measurement. The arithmetic mean and 95% confidence interval for the 13 were 1977 TEEPLE: HERRING GULLS, LAKE ONTARIO ISS TABLE 5—Residue levels of DDE and PCBs in Herring Gull eggs from eastern Canada! Location Year of Number of Residues in ppm dry weight (Number of colonies) collection eggs Geometric mean and 95% CI. DDE PCBs Lake Huron 1972 5 110 368 (2) 65-185 206-656 Lake Erie 1972 6 48.0 300 (3) 33.0-70.1 244-369 Lake Ontario 1972 16 131 565 (8) 104-164 436-730 Lake Ontario 1973 15 134 420 (Brothers Island) 107-167 332-531 Gulf of St. Lawrence 1972 16 32.1 65.2 (2) 20.4-50.6 47.1-90.3 Bay of Fundy 1972 5 14.1 25.4 (1) 7.58-26.1 15.6-41.2 'From Gilbertson and Reynolds (1974 and personal communication), except Brothers Island eggs, which were from this study. 0.339 mm and 0.327-0.351 mm (Table 6). The regression of shell thickness on concentration of DDE in the egg was calculated for the 12 eggs for which both thickness and residue data were available (Figure 2). The F-test showed that the slope of the first order regression line is highly significant (Fi,10 = 12.1, P< 0.01), and that a second order regression gave no significant improvement in fit. Discussion The breeding success found in the Brothers Island Herring Gull colony in 1973, between 0.06 and 0.18 fledged chicks per breeding pair, is among the lowest reported for any North American Herring Gull population. By contrast, Paynter (1949) reported 0.91 fledged young per breeding pair in New Brunswick, Kadlec and Drury (1968) found 0.8—1.4 for various colonies Y = .38.47 - 00035xX | i loa ese | ow al as 380 @ S S B6OK Y) (Up) LJ S 340- O Jc EK/- = _ 320\= ae WY B00F ; [stra] Lah ee As ne 50 75 100 125 TSO WZ DDE CONCENTRATION (PPM, DRY WEIGHT) FIGURE 2. Shell thickness versus DDE concentration in 12 Herring Gull eggs that did not hatch, Brothers Island, 1973. 154 THE CANADIAN FIELD-NATURALIST Vol. 91 TABLE 6—Shell thicknesses and thickness indices for Herring Gull eggs from eastern North America Shell thickness (mm) Thickness index Percent Percent Year(s) of | Number of Arithmetic mean change from Arithmetic mean change from Location collection eggs! 95% C.I. pre-1947 95% C.1. pre-1947 Great Lakes? Pre-1947 456 (369) 0.375 — 1.73 =_ 0.373-0.377 1.72-1.74 Great Lakes? 1960-1969 190 (151) 0.338 -10 1.55 -10 0.334-0.342 1.53-1.57 Lake Michigan} 1970 10 (10) 0.328 -12 1.49 -14 (Green Bay) 0.307-0.349 1.39-1.59 Lake Ontario4 1972 16 (16) 0.309 -18 1.58 ~9 (8 colonies) 0.298-0.320 1.54-1.62 Lake Ontario‘ 1973 11 (13) 0.339 -10 1.60 -8 (Brothers Is.) 0.327-0.351 1.53-1.67 Atlantic Coast? Pre-1947 598 (513) 0.372 — 1.77 ss 0.370-0.374 1.76-1.78 Atlantic Coast? 1963-1969 41 (41) 0.369 =| 1.68 -5 0.361-0.377 1.64-1.72 'Numbers in parentheses are sample sizes for thickness only. 2From Anderson and Hickey (1972). 3From Faber and Hickey (1973). 4From Gilbertson (1974 and personal communication). 5From this paper. on the New England coast, and Keith (1966) reported 0.3-0.4 for a Lake Michigan colony. Reproductive success on Brothers Island was_ similar to that reported by Gilbertson (1974), 0.06—-0.21, for five Lake Ontario colonies. When I first arrived on Brothers Island it was too late to determine the date of onset of nest building, but breeding appeared to be under way normally. Temporal dispersal of first clutch initiations was apparently normal, with 68% of starts taking place during a 12-day interval. Paludan (1951) found 68% of first clutches were initiated during a 14-day interval in both years of his study. In contrast to the prolonged renesting period, as late as mid-July, reported for Herring Gulls on Scotch Bonnet Island in eastern Lake Ontario in 1973 (Gilbertson and Hale 1974), renesting on Brothers Island was completed by 29 May. The primary cause of reduced breeding suc- cess in the Brothers Island colony lay in the failure of 77% of the eggs to hatch. That rate of failure was somewhat higher than the 54-59% reported by Keith (1966) in Lake Michigan, but similar to the 84% reported for Scotch Bonnet Island by Gilbertson and Hale (1974). Among the 37 eggs known to have been infertile or to have suffered embryonic mortal- ity, 33 were incubated, intact, for at least 27 days. Among the 61 eggs that were incubated full term intact (33 that did not hatch and 28 that did) 54% failed to hatch. I believe those 61 eggs constitut- ed an unbiased sample of all the eggs laid in the colony, and indicated that about 54% of all eggs laid were infertile or would have contained dead embryos prior to hatching. Paynter (1949) found that Herring Gull eggs that were infertile or contained dead embryos constituted only 8% of all eggs laid, and Paludan (1951) reported only 6%. The mean number of eggs found in nests at the start of incubation was 2.71 for first clutches and 2.46 for second ciutches. Corresponding clutch sizes reported by Paludan (1951) were 2.95 and 2.77, although it is not entirely clear whether his clutch sizes referred to eggs in nests at start of incubation or to total eggs laid, 1.e., included eggs lost during egg-laying. If eggs lost during egg-laying are included for the Brothers Island colony, the mean sizes for first and second clutches that survived to start of incubation are 2.87 and 2.46, respectively. 1977 Hatching success of eggs laid in 34 first clutches and 13 second clutches was 23% and 22%. In terms of chicks hatched per clutch, first clutches produced 0.62 chicks and second clutches, 0.54. The higher rate for first clutches is primarily due to the higher proportion contain- ing three eggs rather than two. Since the colony was small and the primary object of the study was to determine breeding success, I avoided random collecting of eggs or chicks for studies of embryonic death or chemi- cal residues. But 5 of the 16 eggs collected may be regarded as randomly selected eggs; four, from four clutches, had rolled from the nest or were from a destroyed nest, and one was found, apparently unincubated, where there were no nests. These five are among those for which thickness and thickness index data are presented in Table 6. The remaining I! eggs, from 9 clutches, were selected because they were infer- tile or contained dead embryos. Since Keith (1966) found no significant difference between DDE levels in live and dead Herring Gull eggs, and since mean DDE and PCB levels are not significantly different between eggs of this study and those collected randomly by Gilbertson from eight Lake Ontario colonies in 1972 (Gilbertson and Reynolds 1974, and personal communication) (Table 5), I have regarded those 11 eggs as comparable to randomly collected eggs used in other studies. Kreitzer (1972) reported that complete embryonic development of Coturnix Quail (Coturnix japonica) eggs resulted in a 7.3% thinning of the shells; only two of the Brothers Island gull eggs used for thickness and thickness index determinations showed embryonic development greater than 25%, and seven showed no development what- soever. Shells of the 13 eggs from Brothers Island (Table 6) were significantly thicker (P< 0.001) than the 16 shells collected from Lake Ontario in 1972 by Gilbertson (1974, and personal com- munication). However, they did not differ sig- nificantly in thickness (P > 0.05) from the 1960- 1969 shells on which Anderson and Hickey (1972) reported, nor from the 1970 shells re- ported by Faber and Hickey (1973), and were significantly thinner (P< 0.05) than 369 pre- 1947 shells measured by Anderson and Hickey (1972). Thickness indices (Table 6) were similar TEEPLE: HERRING GULLS, LAKE ONTARIO }15)3) to post-1947 ones, but significantly below those reported for pre-1947 shells (P< 0.05). From the examination of 12 eggs that were incubated but failed to hatch, I could conclude only that embryonic death did not appear to be more common at any particular stage of development. This agrees with Keith’s (1966) findings in a heavily organochlorine-contaminated population, and is at variance with those of Paludan (1951), who found that 80% of eggs that were infertile or had suffered embryonic mor- tality showed no apparent development. Palu- dan’s results probably represent gull populations much less contaminated by organochlorines, since his work was done on the Danish coast in 1943 and 1944. As well as a regression line of eggshell thickness on concentration of DDE in the egg (Figure 2), a regression line of egg-shell thick- ness on concentration of PCBs in the egg (dry weight basis) was also calculated (vy = 0.3834 — 0.00011x, r =0.745). The linearity and the r value are both highly significant, as they are for the DDE regression. This correlation, however, is presumably a spurious one, resulting from the linear relationship known to exist between levels of DDE and PCBs in Great Lakes Herring Gull eggs (Gilbertson 1974). There are several labora- tory feeding studies showing egg-shell thinning effects of sublethal levels of DDE (Cooke 1973), but no such evidence that sublethal levels of PCBs cause any changes in egg-shell thickness (Peakall and Peakall 1973). No obvious cause of death could be deter- mined for 11 of the 12 chicks found dead and collected. Necropsy showed that fecal impaction in the cloaca, possibly related to malnutrition, could have accounted for the death of the other chick. Examination disclosed various disorders, including diarrhoea and malnutrition, in seven chicks, but gave no indication of what could have caused those conditions. Comparison of levels of DDE and DDT in the brains of the Brothers Island chicks with levels found by Hickey et al. (1966) in the brains of adult Herring Gulls indicated that those chemicals were pro- bably not directly responsible for the deaths of the chicks. These comparisons with findings of Hickey et al. (1966) depend on the assumption that lethal brain levels of organochlorines in 156 chicks do not differ greatly from those in adults of the same species. The literature concerning avian brain residue levels of dieldrin, heptachlor epoxide, hexa- chlorobenzene, and PCBs (Jefferies and Prestt 1966; Greenberg and Heye 1971; Faber et al. 1972; Koeman et al. 1973; Stickel et al. 1969) all deals with species other than gulls, and for heptachlor epoxide, hexachlorobenzene, and PCBs, gives no clear indication whether the levels found in the brains of the 12 chicks would have resulted in their deaths. Stickel et al. (1969) concluded that minimum lethal brain residue levels of dieldrin are remarkably similar among bird species, being 4 or 5 ppm wet weight. The highest dieldrin level found in brain tissue of the Brothers Island chicks was 0.47 ppm wet weight, and thus it is unlikely that any of those chicks died from direct dieldrin poisoning. There remains the possibility that the cumula- tive effect of several OC chemicals together was lethal in those chicks that died soon after hatch- ing, before body concentrations of OCs could be reduced by growth. Another possibility is that the excessive egg and chick mortality was due to ineffective parental care by adults carrying high loads of organochlorines. The presence of signs of malnutrition in three of the chicks found dead lends support to this hypothesis. That incuba- tion behavior of birds can be upset by PCBs has been shown experimentally for Ring Doves (Streptopelia risoria) (Peakall and Peakall 1973). I saw people on the island only twice, and I have no evidence to indicate that such visitors, or the fishermen who sometimes anchored off- shore, had any significant effect on breeding success of the gulls. The high pollutant levels and low breeding success found in the Brothers Island Herring Gull colony fit the pattern established for Herring Gulls elsewhere on the Great Lakes. Investigators have attributed other such low success rates to organochlorine pollutants, and the evidence from Brothers Island indicates the same association of high organochlorine levels with low breeding success there. The low breeding success took the form of reduced survival of both eggs and chicks. Whether that reduced survival was related to pollutants in the eggs and chicks, or to pollutants in the adult THE CANADIAN FIELD-NATURALIST Vol. 91 gulls (causing aberrant behavior), or to both, remains unclear. However, the general associa- tion with pollutants is strong, particularly since other possible factors, such as disease, distur- bance, and storms have been evaluated and do not acount for the drastically low breeding success found. Acknowledgments I thank J. A. Keith for the invaluable direc- tion and advice he provided, and R. Hale and I. M. Price for their generous assistance in the field. Others who provided field assistance on several occasions, and to whom I am grateful, were C. G. Gruchy and L. E. Johnston. I extend thanks also to E. Broughton for pathological examination of the specimens, to L. M. Rey- nolds for the chemical analyses, to G. E. J. Smith for advice on the statistical analysis of the data, and to G. Fox, C.G. Gruchy, D. B. Peakall, and P. A. Pearce for critical review of the manuscript. Literature Cited Anderson, D. W. and J. J. Hickey. 1972. Eggshell changes in certain North American birds. /n Proceedings of the XVth International Ornithological Congress. Edited by K. H. Voous. E. J. Brill, Leiden, The Netherlands. pp. 514-540. Cooke, A. S. 1973. Shell thinning in avian eggs by environ- mental pollutants. Environmental Pollution 4(2): 85-152. Faber, A. R. and J.J. Hickey. 1973. Eggshell thinning, chlorinated hydrocarbons, and mercury in inland aquatic bird eggs, 1969 and 1970. Pesticides Monitoring Journal 7(1): 27-36. Faber, A. R., R. W. Risebrough, and H. M. Pratt. 1972. Organochlorines and mercury in Common Egrets and Great Blue Herons. Environmental Pollution 3(2): 111-122. Fimreite, N. and L. M. Reynolds. 1973. Mercury contami- nation of fish in north-western Ontario. Journal of Wildlife Management 37(1): 62-68. Gilbertson, M. 1974. Pollutants in breeding Herring Gulls in the lower Great Lakes. Canadian Field-Naturalist 88(3): 273-280. Gilbertson, M. and R. Hale. 1974. Characteristics of the breeding failure of a colony of Herring Gulls on Lake Ontario. Canadian Field-Naturalist 88(3): 356-358. Gilbertson, M. and L. Reynolds. 1974. DDE and PCB in Canadian birds, 1969 to 1972. Canadian Wildlife Service Occasional Paper Number 19. 18 pp. Goethe, F. 1937. Beobachtungen und Untersuchungen zur Biologie der Silbermowe (Larus a. argentatus Pontopp.) auf der Vogelinsel Memmertsand. Journal fur Orni- thologie 85: 1-119. (Cited in Paludan 1951). Greenberg, R. E.and P. L. Heye. 1971. Insecticide residues in Little Blue Herons. Wilson Bulletin 83(1): 95-97. 1977 Hickey, J.J. and D.W. Anderson. 1968. Chlorinated hydrocarbons and eggshell changes in raptorial and fish- eating birds. Science (Washington) 162(3850): 271-273. Hickey, J. J., J. A. Keith, and F. B. Coon. 1966. An ex- ploration of pesticides in a Lake Michigan ecosystem. Journal of Applied Ecology 3(Supplement): 141-154. Jefferies, D. J. and I. Prestt. 1966. Post-mortems of Pere- grines and Lanners with particular reference to organo- chlorine residues. British Birds 59(2): 49-64. Kadlec, J. A. and W.H. Drury. 1968. Structure of the New England Herring Gull population. Ecology 49(4): 644-676. Keith, J. A. 1966. Reproduction in a population of Herring Gulls (Larus argentatus) contaminated by DDT. Journal of Applied Ecology 3(Supplement): 57-70. Koeman, J.H., H.C. W. van Velzen-Blad, R. de Vries, and J.G. Vos. 1973. Effects of PCB and DDE in cor- morants and evaluation of PCB residues from an experi- mental study. Journal of Reproduction and Fertility, Supplement 19: 353-364. Kreitzer, J. F. 1972. The effect of embryonic development on the thickness of the egg shells of Coturnix Quail. Poultry Science 51(5): 1764, 1765. Ludwig, J. P. and C.S. Tomoff. 1966. Reproductive suc- cess and insecticide residues in Lake Michigan Herring Gulls. Jack-Pine Warbler 44(2): 77-85. TEEPLE: HERRING GULLS, LAKE ONTARIO 157 Paludan, K. 1951. Contributions to the breeding biology of Larus argentatus and Larus fuscus. Ejnar Munksgaard, 6, Norregade, Copenhagen. 142 pp. Paynter, R. A., Jr. 1949. Clutch-size and the egg and chick mortality of Kent Island Herring Gulls. Ecology 30(2): 146-166. Peakall, D. B. and M. L. Peakall. 1973. Effect of a poly- chlorinated bipheny] on the reproduction of artificially and naturally incubated dove eggs. Journal of Applied Ecology 10(3): 863-868. Ratcliffe, D. A. 1967. Decrease in eggshell weight in certain birds of prey. Nature (London) 215(5097): 208-210. Reynolds, L. M. and T. Cooper. 1975. Analysis of organo- chlorine residues in fish. Jn Water Quality Parameters, ASTM STP 573. American Society for Testing and Materials. pp. 196-205. Stickel, W.H., L.F. Stickel, and J.W. Spann. 1969. Tissue residues of dieldrin in relation to mortality in birds and mammals. /n Chemical fallout; current research on persistent pesticides. Proceedings of the First Rochester Conference on Toxicity. Edited by M. W. Miller and G. G. Berg. Charles C. Thomas, Springfield, Illinois. pp. 174-204. Received 5 January 1976 Accepted 10 February 1977 Weedy Species at Terminal Grain Elevators, Thunder Bay, Ontario DOUGLAS R. LINDSAY Department of Biology, Lakehead University, Thunder Bay, Ontario Lindsay, Douglas R. 1977. 91(2): 158-164. P7B 5El Weedy species at terminal grain elevators, Thunder Bay, Ontario. Canadian Field-Naturalist Abstract. In 1975, a survey of adventive plants was carried out at the terminal grain elevators and adjacent railway yards, Thunder Bay, Ontario. A total of 146 species was recorded, of which 27 species have likely been introduced as contaminants in western grain shipments. Eleven species common to eastern Canada were also recorded. The weed flora at Thunder Bay shows somewhat similar composition to that found at another grain terminal at Churchill, Manitoba. There is no evidence that these adventives pose any serious threat to local agriculture. The 19 terminal grain elevators at Thunder Bay, Ontario afford an unusual opportunity for study of ruderal assemblages of plants trans- ported, in some cases, long distances as con- taminants of grain shipments. Grain spillage from box cars is rather common and such spills may inadvertently lead to the introduction of weeds which have been previously harvested with associated grain or other crops. An earlier survey by Beckett (1959) lists 76 species of adventives at the terminal elevator and railway yards of Churchill, Manitoba. She reports that 28 species probably do not persist but are introduced sporadically in grain ship- ments to that port. Unfortunately, her work has not been repeated so that an up-to-date listing of persistent species is not available. In general, railway yards and rights-of-way provide unusual habitats for weedy plants. Such habitats differ considerably from those of cul- tivated fields, pastures, rangeland, and even vacant lots. In Wisconsin, Curtis (1959) showed a number of weeds achieved greater presence values in railway yards than in other plant communities. Railway yards must have adequate drainage. Thus track ballast must be largely course aggre- gates of gravel and sand. In summer, heat and moisture conditions may fluctuate greatly and in winter snow cover is generally minimal. Even highway road shoulders, where some conditions may be similar to those of railway ballast, undergo more frequent grading and accumula- tions of salt than is found in railway yards. Soil nutrient conditions may also be important. The purpose of the present study was three- 158 fold. Firstly, a check-list of all adventives was made at the terminal elevators. Secondly, this flora was compared with the studies carried out by Beckett (1959) and Curtis (1959) and finally, a check of local grain fields was carried out to ascertain if the grain terminal weeds posed any threat to local agriculture. Description of Study Area Figure | shows the general location of the area surveyed. For convenience, it was divided into eight sites commencing with the most westerly portion known as the “Neebing Yard” (Site 1) and ends with site 8 in the northeast section of Thunder Bay. The total distance between sites | and 8 is about 20 k (12.6 mi). A typical site is shown in Figure 2. The over-all terrain is generally flat and was formerly submerged by high levels of Lake Superior during the last post-glacial period. The original organic soils have been greatly modified by improved drainage and track ballast in the railway yards. For the most part, all sites except site 5 are rather uniform as to age of existence. Site 5, known as the Keefer Terminal, was opened in 1962 for shipment of general cargo and is not involved in the trans shipment of grain as are the elevators. Also, site 7 is a small holding stockyard for livestock and was included in the survey because of possible weed introduction in hay used to feed the animals. Methods Each site was visited several times from May through September 1975 and presence lists were compiled for each site. Allintroduced weeds and 1977 LINDSAY: WEEDS AT GRAIN ELEVATORS, THUNDER BAY 159 FiGurE 1. Location of eight sites sampled at terminal grain elevators and railway yards, Thunder Bay, Ontario. native adventive plants were recorded and voucher specimens collected. The area surveyed included 17 of 19 elevators with their adjacent yards. Intervening track areas between sites were also traversed on foot. Although some areas were restricted, and, owing to hazards of shunt- ing trains, all tracks could not be surveyed; I am confident, however, that few plants escaped detection. Percentage of species in each plant family was calculated and compared with results obtained by Beckett (1959) and Curtis (1959). Species common to all three locations, Churchill, Wis- consin, and Thunder Bay, were also determined. A total of 12 grain fields in the Lakehead area was surveyed for possible introduction of weeds found at the elevators. 160 THE CANADIAN FIELD-NATURALIST Vol. 91 FIGURE 2. Typical site (site 8). Toadflax (Linaria vulgaris) in foreground. This elevator was demolished in 1976. Weeds of Canada (Frankton and Mulligan 1970) was used as a guide for the study. Results Table | lists by family all adventive plants found in this study. Species predominantly weedy in western Canada are preceded by ‘w’ and those of eastern Canada by ‘e’. Autogamous species, previously determined by Mulligan and Findlay (1970), are designated by ‘o’. folium, Weeds found at all sites included Equisetum arvense, Agropyron repens, Panicum capillare var. capillare, Polygonum aviculare, Cheno- podium album, Salsola pestifer, Amaranthus retroflexus, Capsella bursa-pastoris, Lepidium densiflorum, Potentilla norvegica, Medicago lupulina, Melilotus alba, Trifolium pratense, Linaria vulgaris, Plantago major, Achillea mille- Artemisia biennis, Crepis tectorum, Erigeron canadensis, Matricaria maritima var. 1977 LINDSAY: WEEDS AT GRAIN ELEVATORS, THUNDER BAY 161 TABLE |—List of adventive plants compiled from all sites. w, Weedy in the Prairie Provinces; e, weedy in eastern Canada; 0, autogamous species (after Mulligan and Findlay 1970) Equisetaceae Caryophyllaceae (cont.) Equisteum arvense L. w Silene cserei Baumg. E. hymenale L. var. affine (Englm.) A. A. Eat. Con Sci cuba seWabel Gramineae o S. noctiflora L. Agropyron repens (L.) Beauv. o Stellaria media (L.) Vill. Agrostis scabra Willd. w Avena fatua L. Beckmannia syzigachne (Steud.) Fern. Ranunculaceae eo Ranunculus acris L. Bromus inermis Leyss. Papaveraceae 3 e Digitaria ischaemum (Schreb.) Muhl. Corydalis aurea Willd. w Echinocloa crusgalli (L.) Beauv. (G@ruciterac w Hordeum jubatum L. Arabis glabra (L.) Bernh. w Lolium persicum Boiss. & Hoh. Armoracia lapathifolia Gilib. e Panicum capillare L. var capillare : e Barbarea vulgaris R. Br. Brassica campetris L. B. juncea (L.) Czern. 0 Capsella bursa-pastoris (L.) Medic. wo Descurainia sophia (L.) Webb. w Erucastrum gallicum (Willd.) O. E. Schulz o Erysimum cheiranthoides L. Urticaceae Lepidium campestre (L.) R. Br. Urtica dioica L. var. procera Wedd. o L. densiflorum Schrad. Sinapis alba L. wo S. arvensis L. wo Sisymbrium altissimum L. P. miliaceum L. Phleum pratense L. Poa compressa L. P. pratensis L. Puccinellia nuttalliana (Schultes) Hitch. wo Setaria viridis (L.) Beauv. Santalaceae Commanadra richardsiana Fern. Polygonaceae w S. loeselti Ik Rumex acetosella L. wo Thlaspi arvense IL R. maritimus L. var. fueginus (Phillipi) Dusen. o Rorippa islandica (Oeder) Borbas R. mexicanus Meisn. : Saxifragaceae w R. pseudonatronatus Borbas pa oL F Be eienophylius Ledeb: ssia palustris L. var. neogaea Fern. Polygonum achoreum Blake Rosaceae o P. aviculare L. o Potentilla argentea L. o P. convolvulus L. P. anserina L. o P. lapathifolum L. P. norvegica L. o P. persicaria L. i P. ramossisimum Michx. Leguminosae : BPE conriumiocnch: Lotus corniculatus L. 0 Medicago lupulina L. Chenopodiaceae M. sativa L. Atriplex hortensis L. Melilotus alba Dest. A. patula L. var. hastata (L.) Gray M. officinalis (L.) Lam. w Axpris amaranthoides L. Trifolium hybridum L. ° Chenopodium album L. C. glaucum L. C. gigantospermum Aellen w Corispermum hyssopifolium L. T. pratense L. T. procumbens L. Vicia cracca L. ° w Kochia scoparia (L.) Schrad. Euphorbiaceae w Salsola pestifer Nels Euphorbia glyptosperma Engelm. Amaranthaceae Malvaceae Amaranthus albus L. Malva pusilla Sm. A. blitoides S. Wats. Onagraceae i xus L. WO) Als aliiaizee) Epilobium angustifolium L. Caryophyllaceae E. glandulosum Lehm. var. adenocaulon (Haussk.) 0 Cerastium vulgatum L. Fern. o Lychnis alba Mill. 0 Oenothera biennis L. 162 Table I (continued) THE CANADIAN FIELD-NATURALIST Vol. 91 Umbelliferae Pastinaca sativa L. Apocynaceae Apocynum androsaemifolium L. Asclepiadaceae e Asclepias syriaca L. Polemoniaceae Collomia linearis Nutt. Boraginaceae eo Echium vulgare L. w Lappula echinata Gilib. Labiatae w Galeopsis tetrahit L. var bifida (Boenn.) Lej. & Court. Mentha arvensis L. Scrophulariaceae Chenorrhinum minus (L.) Lange. Euphrasia hudsoniana Fern. & Weig. Linaria vulgaris Mill. Odontites serotina (Lam.) Dum. eo Verbascum thapsus L. Plantaginaceae o Plantago major L. Rubiaceae Galium boreale L. Compositae Achillea millefolium L. e Ambrosia artemistifolia L. var elatior (L.) Desc. A. psilostachya DC. var. coronopifolia (T. & G.) Farw. e A. trifida L. Anaphalis margaritacea (L.) C. B. Clarke 0 Arctium minus (Hill) Bernh. wo Artemisia biennis Willd. A. caudata Michx. A. absinthium L. A. ludoviciana Nutt. var. gnaphalodes (Nutt.) Compositae (cont.) T. & G. A. vulgaris L. Aster ericoides L. A. modestus Lindl. A. prealtus L. A. simplex Willd. var. ramossisimus (T. & G.) Cronq. Bidens cernua L. B. frondosa L. Chrysanthemum leucanthemum L. var. pinnatifi- dum Lecog & Lamotte e Cichorium intybus L. Cirsium arvense (L.) Scop. C. vulgare (Savi) Tenore Crepis tectorum L. Erigeron (Conyza) canadensis L. E. strigosus Muhl. Grindelia squarrosa (Pursh) Dunal Helianthus annuus L. H. maximilliana Schrad. H. laetiflorus Pers. var. subrhomboideus (Rydb.) Fern. H. tuberosus L. Hieracium canadense Michx. H. floribundum Wimm. & Grab. w Iva xanthifolia Nutt. Matricaria maritima L. var. agrestis (Knaf) Wilmott o M. matricarioides (Less.) Porter Rudbeckia serotina Nutt. o Senecio vulgaris L. Solidago canadensis L. S. gigantea Ait. S. graminifolia (L.) Salisb. w Sonchus arvensis L. var. glabrescens Guenth., Grab., & Wimm. Tanacetum vulgare L. 0 Taraxacum officinale Weber wo Tragopogon dubius Scop. Xanthium strumarium L. ZEO0O0 ZO agrestis, M. matricariodes, Solidago canadensis, Sonchus arvensis var. glabrescens, and Taraxa- cum officinale. Some 55 species are recorded for the more recent Keefer Terminal (site 5), including the native plants Parnassia palustris var. neogaea, Euphrasia hudsoniana, and Odontites serotina. Such plants are probably “hold-overs” from a period prior to 1962 when the terminal was opened. This site has not yet acquired all of the species found at the elevators. At the stockyard (site 7) the only survey record for Barbarea vulgaris was found. This plant was likely introduced in hay. Some of the most serious weeds of western Canada such as Euphorbia esula L., Cardaria draba (L.) Desv., Neslia paniculata (L.) Desv., Camelina microcarpa Andrz., Conringia orien- talis (L.) Dumort., Dracocephalum parviflorum Nutt., /va axillaris Pursh, and Lactuca pul- chella (Pursh) DC. were not found. Records for all the preceding except Cardaria draba are known for the Thunder Bay District. The present survey also shows a possible new record for the province, Rumex stenophyllus Ledeb. which has been introduced from western Canada. Of the eastern weeds, Daucus carota L., 1977 Convolvulus arvensis L., Hypericum perfora- tum L., and Spergula arvensis L. were not found but records are known for these plants in the district. Table 2 shows a comparison of the principal plant families found at three widely separated locations. The high value for Compositae at Thunder Bay is due in part to the large number of native plants found in this study. Forty-nine species found at Thunder Bay were also found in the Churchill study and 46 species were similar to a weed list based on data from 24 railway yards in Wisconsin. Species common to all three locations include Poa pratensis, Agro- Pyron repens, Hordeum jubatum, Phleum pra- tense, Setaria viridis, Polygonum aviculare, P. convolvulus, Chenopodium album, C. glaucum, C. gigantospermum, Lychnis alba, Lepidium densiflorum, Erucastrum gallicum, Potentilla norvegica, Trifolium hybridum, Melilotus alba, M. officinalis, Collomia linearis, Linaria vul- garis, Plantago major, Iva xanthifolia, Arte- misia biennis, Cirsium arvense, and Taraxacum officinale. In the 12 local grain fields surveyed only Sonchus arvensis var. glabrescens and Cheno- podium album were found in all fields, Cirsium arvense in four, Arena fatua in two, and Poly- gonum scabrum and Setaria viridis in one each. The “eastern” Spergula arvensis was found on one field but not at the elevator sites. Discussion and Conclusions There is little doubt that many weed species are introduced at the terminal elevators of Thunder Bay through leakage and grain spills from railway box cars. Unfortunately, I was not LINDSAY: WEEDS AT GRAIN ELEVATORS, THUNDER BAY 163 able to analyze grain samples for identification of weed contaminants and to correlate species with those found on the sites. The new Govern- ment of Canada hopper cars minimize such spillage and if some further cleaning could be done at points of origin, further reduction of contaminants would result. A few elevators have landscaped their property and this has resulted in more pleasing aesthetic effect. Spraying of yard species, in my view, would only increase a fire hazard. This study has not been able to prove per- sistence of these weeds. It is suspected that some plants such as Arena fatua, Lolium persicum, and Axyris amaranthoides must be introduced each year. Others such as Linaria vulgaris and Crepis tectorum appear to be well established and have spread into suitable habitats within the city. Ragweeds (Ambrosia spp., Iva spp.) and the members of the Chenopodiaceae pose some threat to hay-fever sufferers. Thus far, however, the elevator plants are not a menace to local grain fields which are rather scattered and of small acreage. Floristic comparisons between the present study and other railway yards have shown a similarity, in which the major component of each is encompassed by a few plant families. At Thunder Bay, some 38 species are autogamous weeds, as determined by Mulligan and Findlay (1970). Mulligan (1972) has pointed out that such plants may have a decided advantage for establishment after long-distance dispersal. Of the 24 species common to all three railway yard locations, nine are autogamous species. Re- cently, Grime (1974) has used a triangular ordination of vegetation in the Sheffield area in TABLE 2—Percentage of total species in principal plant families represented at three railway yard locations Plant Curtis family (Wisconsin) Compositae 18.6 Gramineae 14.8 Cruciferae Wed Polygonaceae 6.6 Leguminosae 4.9 not recorded not recorded Chenopodiaceae Caryophyllaceae Beckett Present study (Churchill) (Thunder Bay) 13.2 30.2 15.8 10.9 21.0 11.6 5.2 8.2 10.5 6.1 10.5 6.1 Se 4.1 164 Great Britain, based on degree of competition, stress, and disturbance. He has shown ordina- tion patterns of railway ballast plants to be similar to some communities of wasteland but quite different from those of meadows, pastures, and roadsides. It would seem that further studies would be useful to develop more quantitative methods in weed ecology. Lastly, more work should be done on the environmental factors found in railway yards. In this study Beckmannia syzigachne and Puccinellia nutalliana are native plants found in more saline soils of western Canada. The physical and chemical properties of ballast soils, moisture and temperature regimes as related to the life cycles of these and other plants might be revealing. Acknowledgments I am grateful to the National Research Council of Canada (Grant A9880) for financial THE CANADIAN FIELD-NATURALIST Vol. 91 assistance. Sincere appreciation and thanks is given W.R. Russell, Department of Biology, Lakehead University for his valuable assistance in the survey. Literature Cited Beckett, E. 1959. Adventive plants at Churchill, Manitoba. Canadian Field-Naturalist 73: 169-173. Curtis, J. T. 1959. An ordination of plant communities. Jn The vegetation of Wisconsin. University of Wisconsin Press, Madison, Wisconsin. pp. 412-434, 591-596. Frankton, C. and G. A. Mulligan. 1970. Weeds of Canada. Canada Department of Agriculture Publication 948. Grime, J. P. 1974. Vegetation classification by reference to strategies. Nature (London) 250: 26-31. Mulligan, G. A. 1972. Autogamy, allogamy and pollina- tion in some Canadian weeds. Canadian Journal of Botany 50: 1767-1771. Mulligan, G. A. and J. Findlay. 1970. Reproductive sys- tems and colonization in Canadian weeds. Canadian Journal of Botany 48: 859-860. Received 11 February 1976 Accepted 10 February 1977 Increase in Overwintering by the American Goldfinch, Carduelis tristis, in Ontario A. L. A. MIDDLETON Department of Zoology, University of Guelph, Guelph, Ontario NIG 2WI1 Middleton, A. L. A. 1977. Field-Naturalist 91(2): 165-172. Increase in overwintering by the American Goldfinch, Carduelis tristis, in Ontario. Canadian Abstract. Between mid-November and mid-May from 1970 until 1975, 3433 American Goldfinches, Carduelis tristis, were banded at Guelph, Ontario. Banding results show that many goldfinches now winter in Guelph. Analysis of the annual Christmas Bird Count data shows that a significant increase has occurred in overwintering goldfinch populations in southern Ontario since 1915. This population increase cannot be attributed solely to climatic amelioration, and is apparently related to an abundance of food provided at feeding stations. As a result the goldfinch is now a common winter resident in the urban areas of southern Ontario. Snyder (1957) and de Vos (1964) have sum- marized the marked changes which have occur- red in the avifauna of Ontario during the past century, while von Haartman (1973) has re- ported similar changes in northern Europe. Three causes have been proposed to explain these changes: an increase in the number of bird watchers, an amelioration of the climate, and an alteration of the habitat by man (von Haartman 1973). In addition to the obvious changes which accompany habitat alteration, less obvious and more subtle changes, such as the adaptation by indigenous species to urban environments and conditions, have occurred and probably still are occurring (Tast 1968; Ward 1968). In Ontario the number of bird watchers has undoubtedly increased (records of annual Christmas Bird Counts), the climate is probably a little milder than a century ago, although the last 20 years has seen a return to colder, snowier conditions (M. K. Thomas, unpublished manu- script), and modern agriculture and urbaniza- tion have modified the original environments and created new ones. In response to man-made changes, birds either decline (Wallace 1970; Batten 1972; Emlen 1974) or adapt to the new environment (Tast 1968; Ward 1968; Boshko 1971; Emlen 1974). In those parts of the world recently settled by European man, e.g., North America and Australia, adaptation by birds to ecologically disturbed conditions is presumably at an earlier stage than in Europe. Thus many avian species in North America such as the Blue Jay, Cyanocitta cristata, (Bock and Lepthien 1976a) may be adapting to new conditions. In Guelph, Ontario, a large population of the American Goldfinch, Carduelis tristis, now regularly overwinters, whereas wintering by goldfinches in Ontario was not common in former years (Mcllwraith 1894; Snyder 1951). This paper documents an increase in winter populations of the American Goldfinch in Ontario during the past 60 years and attempts to explain its cause. Methods and Materials Between mid-November and mid-May 1970-1975, goldfinches were trapped and banded at Guelph, Ontario using baited Potter traps. At capture each bird was sexed and weighed to the nearest 0.5 g. For the purposes of this paper, banding data were used only to determine the size of the overwintering popula- tions at Guelph (Davis 1963). Although banding was mainly carried out at two locations, sepa- rated by 1.5 km in thecenter and west of the city, four other sites were used during the study (all to the south of the main areas, within a radius of 4.5 km). In addition my banded birds were captured by two other banders within Guelph. Data on winter goldfinch populations in southern Ontario (localities south of 46° N) were obtained from the published results of the annual Christmas Bird Counts. The data were calculated on the basis of number of birds counted per party hour per year. This method was considered the most appropriate (Raynor 1975) and valid in view of the various sources of error inherent in the original data (Bock and Smith 1971). 165 166 In a part of Ontario as large as that defined above it is difficult to obtain weather data which are equally applicable to the entire area. But, accepting that an intensive study of few data from very few representative locations gives a valid indication of weather trends over a large area (Thomas, personal communication) and that weather data from a single station in southern Ontario roughly parallel those from other stations in the same area (Thomas 1957), it was felt that the Guelph data would suffice to show climatic trends throughout southern Ontario since 1915. Thus, 10-year moving means (Thomas 1968) were calculated for temperature and snowfall in Guelph in December (the month in which most Christmas Bird Counts are made), from data provided by the Weather Records Branch, Department of Land Resource Science, University of Guelph. These data were supple- mented by data from Environment Canada for temperature in Ontario and snowfall in Ontario and Quebec (Thomas 1975). In the absence of matching long-term Christmas Count data from Guelph, a regression analysis was made between the weather data and the goldfinch data for Ontario to see whether any correlation existed. An attempt was made to determine the size and importance of the wild-bird seed industry within Ontario and the scale on which wild-bird feeding has increased in recent years. Statistics on the sales of wild-bird seed in the province were sought from 25 companies, both wholesale and retail, and from the Ontario Ministry of Agriculture and Food and the Canadian Seed Trade Association. Results During the study, 3433 goldfinches were THE CANADIAN FIELD-NATURALIST Vol. 91 banded at Guelph, Ontario. Considerable move- ment of banded birds apparently occurred within the city during winter since the per- centage of retraps at the site of last capture varied 46-55% in different years. Between 1970 and 1975, with the exception of the unusually mild and wet autumn of 1974, goldfinches first appeared at feeding stations within Guelph in mid-November. By late November large flocks were usually common within the city. This winter population remained until early May when feeding stations were gradually abandoned by the birds. A simple analysis of the capture-recapture data for April showed that the number of birds in over- wintering flocks in Guelph varied between 853 in 1971-72 and 1816 in 1972-73 (Table 1). Figure | shows the fluctuation in winter weights of goldfinches at Guelph. No significant differences were found in the weights between years. Asa result the goldfinch weight data were pooled by sex. Maximum body weight was reached in December-January, followed by a gradual decrease until mid-March. A slight increase in weight occurred in late March followed by a significant (P< 0.05, Student’s t-test) decrease in April and May. Pre-nuptial molt begins in the population in mid-March and continues throughout April and May (Middle- ton, in press). Migration apparently occurs during May (Tyler 1968), coinciding in Guelph with the abandonment of feeding stations by the winter goldfinch population. Figure 2, from the Christmas Bird Count data for all centers in Ontario, shows an increase in the winter goldfinch populations between 1915 and 1975. The average number of birds counted / party hour increased significantly (P< 0.01, TABLE 1—Population estimates (N = “ty for April goldfinch flocks, Guelph, Ontario, 1970-1975 Total April Number Population 95% Season banded, sample, marked, estimate, confidence M n m N+SE limits 1970-71 541 64 22 1574 + 271.8 1030-2118 1971-72 461 124 67 853 + 70.7 711-995 1972-73 536 166 49 1816 + 217.8 1380-2252 1973-74 751 57 35 1225 + 128.6 967-1483 1974-75 533 96 47 1089 + 113.5 863-1315 1977 17 15 Gone = aie © lu 43 = e MALE o- - - FEMALE 12 MIDDLETON: AMERICAN GOLDFINCH OVERWINTERING IN ONTARIO 167 MONTH FIGURE |. Mean (+ SE) body weight of goldfinches trapped during winter at Guelph, Ontario, 1967-1974. Horizontal bars indicate one standard error; numbers give sample size for each sex. Mann-Whitney U-test) from 0.12 + 0.004 in 1915-1924 to 1.76 + 0.183 in 1966-1975. The 10-year moving means for temperature and snowfall for December at Guelph are shown in Figure 3. December temperatures reached a peak in the mid-1950s, followed by much colder conditions and a recent gradual return to milder temperatures. Snowfall declined steadily until the early 1960s since when there has been a return to much snowier conditions. These data are reflected in the annual data compiled by Environment Canada (Figure 4). Regression analyses showed a significant (P< 0.01) negative correlation between snow- fall and population (Y= 28.525 — 1.713x, r=-0.392, n=60) and a less significant (P< 0.05) negative correlation between temp- erature and population (Y = -3.971 + -119x, r=-0.25, n= 60). Analysis of the data for 50 years (1915-1964) instead of 60 years (1915-1974), however, showed the correlation between snowfall and population to be more pronounced (r = -0.546, P < 0.01) whereas that of temperature was insignificant (r = —0.118). 168 THE CANADIAN FIELD-NATURALIST Vol. 91 15 20 25 30 35 40 45 50 55 60 65 70 75 TIME (YEARS) FIGURE 2. Index of winter American Goldfinch numbers within southern Ontario, 1915-1975, as calculated from annual Christmas Bird Count data. 3] TEMPERATURE MEAN TEMP (°C) 361 SNOWFALL WwW NO MEAN SNOWFALL (CM) nN nN ™ oo Le) (o) 19 20 25 30 35 40 45 350 55 BO OS V0) 7/5 DECADAL YEAR FIGURE 3. Ten-year moving means of snowfall and temperature at Guelph, Ontario, 1915-1974. OH MIDDLETON: AMERICAN GOLDFINCH OVERWINTERING IN ONTARIO 169 TEMPERATURE 50 55 40 45 #£50 55 60 65 70 75 ® Fi O =| 60 65 70 75 ANNUAL SNOWFALL FIGURE 4. Temperature and snowfall trends for southern Canada, 1940-1970, as based on records of Environment Canada (Thomas 1975). (Temperature data from airports at Kapuskasing, London, and Ottawa. Snowfall data from airports at Toronto, Ottawa, Montreal, and Quebec City.) Over the 4-year period for which data were obtained (1969-1973), wholesale sales (records of three companies) of seed for feeding wild birds rose by 50% from 340 tons to 510 tons. Retail sales for the same period (records of five companies) showed an increase of 131% from 350 tons to 810 tons. Discussion Traditionally it has been accepted that gold- finches in Ontario and neighboring regions migrate south for the winter (Dawson 1903; Dionne 1906; Snyder 1951; Godfrey 1966; Tyler 1968). It is known that goldfinches do migrate (records of Bird-Banding Office, United States Fish and Wildlife Service; A. L. A. Middleton, unpublished), but there are records to suggest that some goldfinches have consistently attemp- ted to overwinter in southern Ontario and neigh- boring regions (MclIlwraith 1894; Dawson 1903; Barrows 1912; Snyder 1951; Forbush and May 1955). Attempts at overwintering may have been more common than reported as the goldfinch 1s inconspicuous during winter and often feeds with other species (Tyler 1968). Additionally, as the goldfinch feeds mainly on the seeds of the compositae (A. L. A. Middleton, unpublished) it 1s less dependent on the seeds of trees than other northern carduelines. As a result, its populations do not show the marked fluctua- tions during winter which make other species conspicuous by their presence or absence (Bock and Lepthien 1976b). Thus the presence of the goldfinch can be easily overlooked. The evidence shows, however, that during the last 60 years overwintering by goldfinches in Ontario has increased significantly (Figure 2) and that large numbers of goldfinches now regularly over- 170 winter in Guelph. The winter weights at Guelph roughly corres- pond to those of the goldfinch in Ohio (Wise- man 1975) and show a similar trend to those of other finches in winter (Bartleson and Jenson 1955; King and Farner 1966; Newton 1969, 1972). The increase in weight observed in late March - early April is apparently associated with the prenuptial molt (Middleton, in press). The significant decrease in weight which follows is probably related to migration. An increase in body weight is common in molting birds (King and Farner 1966; Haukioja 1969; Newton 1969; Payne 1972), resulting from an increase in lean dry weight, fat, water content of the feather, and blood volume, all associated with growth of the feather papilla (Payne 1972). But molt also involves increased energy demands and an increase in metabolic rate (Payne 1972). Late March and April in southern Ontario are climatically unpredictable with highly variable temperatures (range of means —6.9° to 9.5°C), irregular snowfalls (range of means -0.0 to 54.9 cm), and frequent freezing rain storms (Weather Records Branch, Department of Land Resource Science, University of Guelph). Such unpredictable and stormy weather is known to cause increased mortality in molting birds (Haukioja 1969), yet at Guelph there was no apparent increase in mortality and the birds gained weight. Newton (1967) has pointed out that when feeding conditions are worst a bird’s food requirements are often greatest. Starving bullfinches, Pyrrhula pyrrhula, could outlive their fat reserves for only a few hours, and Newton (1969) suggested that in hard weather the ability to accumulate sufficient fat each day was a critical factor affecting overnight survival. Thus to survive the Ontario winter, gold- finches have to find sufficient food during January and February, and then to survive the prenuptial molt with its increased energy demands and decreased plumage insulation at a time of inclement weather and food shortage. It seems likely, therefore, that late winter formerly must have been a time of high mortality in any population of goldfinches overwintering in southern Ontario. Winter weather has ameliorated slightly dur- ing the past century, but since the late 1950s mean winter temperatures have been lower, THE CANADIAN FIELD-NATURALIST Vol. 91 accompanied by a return to snowier conditions (M.K. Thomas, unpublished manuscript). These trends are reflected in the December weather conditions at Guelph (Figure 3) and for Ontario as a whole (Figure 4). Within large urban areas, however, an “urban effect” often produces a dome of air over the city which may be several degrees warmer than the air of the surrounding countryside (Thomas 1968). If, as argued, the December weather condi- tions at Guelph refiect the conditions through- out southern Ontario, there is a distinct negative correlation between temperature and snowfall with goldfinch population increases, snowfall having the higher correlation. These data sug- gest that in the early part of the population increase, the goldfinch responded directly to reduced snowfall by attempting to overwinter in Ontario. A decrease in snowfall would have resulted in the prolonged availability of, and easier access to, the natural food supply. At the present time the goldfinch may still respond directly to snowfall since its appearance at winter feeding stations coincides with the first snowfalls in November. The poor correlations between population increase and temperature suggest that temperature in itself has not contributed directly to the observed population increases, although the influence of the urban effect is not known. Thus climatic amelioration, as evidenced by reduced snowfall, probably influenced the increase in goldfinch populations in its early stages, but the evidence now suggests that winter populations are continuing to in- crease, uninfluenced by weather conditions. As von Haartman (1973) has indicated, climatic change alone cannot explain the marked changes in the avifauna of northern Europe, and it seems unlikely that climatic amelioration can be the sole cause of overwintering by goldfinches in Ontario. Among others, de Vos (1964), von Haartman (1973), Alison (1976), and Bock and Lepthien (1976a) have pointed out the importance of winter feeding in the increased overwinter survival and range expansion of several species, and I suspect this is the key to successful over- wintering by goldfinches in Ontario. It is difficult to estimate the real increase in the number of feeding stations in a city such as Guelph during the past, but in Ontario the sales SEF of bird seed have shown a marked growth in recent years, probably continuing a trend esta- blished previously. Although each individual feeder may supply a limited amount of food each day, the total amount of food supplied by all the people feeding wild birds within a city such as Guelph must be considerable. Emlen (1974) estimated that on his study area in Tucson, Arizona, an area little affected by severe winter weather, about half of the avian community’s need was provided by food at feeding stations. The abundance of food now being provided during winter within settled areas of Ontario has created a new situation for some birds. For seed- eating species, winter need no longer be aseason of food shortage, provided these species can exploit the new food supply. The regularity with which the goldfinch reappears at feeding stations in Guelph, coincident with the first snows of winter, suggests that this species now recognizes the urban environment as a reliable food source when conditions are severe. The amount of movement which occurs between feeding sta- tions suggests the birds move readily through- out the city, stopping wherever food can be found. There is no reason to believe that gold- finches in Guelph should behave differently from those in other parts of southern Ontario. Thus, with its ability to exploit this new, abundant food supply, the goldfinch now has become a common winter resident in the urbanized areas of southern Ontario. Acknowledgments I gratefully acknowledge the assistance of Mary Gartshore, Mike Dyer, Murray Pengelley, and Alex Derry who assisted in the banding programs. I thank Robert Prys-Jones and lan Newton for their critical comments and dis- cussion on earlier drafts of this paper. Thanks are expressed to those companies which res- ponded to my request for information on sales of bird seed; to M. K. Thomas for his assistance with, and permission to use unpublished cli- matic data; and tothe R. H. Manskes of Guelph, who permitted me to carry out banding at their feeding station and in the comfort of their home. Literature Cited Alison, R. M. 1976. Mourning Doves wintering in Ontario. Canadian Field-Naturalist 90: 174-176. MIDDLETON: AMERICAN GOLDFINCH OVERWINTERING IN ONTARIO hl Barrows, W.B. 1912. Michigan bird life. Department of Zoology and Physiology, Michigan Agricultural College, Special Bulletin. 822 pp. Bartleson, F.D., Jr. and O. F. Jensen. 1955. A study of Purple Finch winter weights. Wilson Bulletin 67: 55-59. Batten, L.A. 1972. Breeding bird species diversity in relation to increasing urbanisation. Bird Study 19: 157-166. Bock, C. E. and L. W. Lepthien. 1976a. Changing winter distribution and abundance of the Blue Jay, 1962-1971. American Midland Naturalist 96: 232-235. Bock, C.E. and L.W. Lepthien. 1976b. Synchronous eruptions of boreal seed-eating birds. American Naturalist 110: 559-571. Bock, C. E. and R. B. Smith. 1971. An analysis of Col- orado Christmas Counts. American Birds 25: 945-947. Boshko, S. 1971. The process characteristics of avian urbanization. Leningradski Universitet Vestnick 9: 5-14. Davis, D. C. 1963. Estimating the numbers of game pop- ulations. /n Wildlife investigational techniques. Second edition. Edited by H. S. Mosby. Chapter 5. The Wildlife Society, Ann Arbor. Dawson, W.L. 1903. The birds of Ohio. Volume I. Wheaton Publishing Company, Columbus, Ohio. 368 pp. de Vos, A. 1964. Range changes of birds in the Great Lakes Region. American Midland Naturalist 71: 489-502. Dionne, C. E. 1906. Les oiseaux de la province de Québec. Dussaultet Proulx, Quebec. 414 pp. Emlen, J. T. 1974. An urban bird community in Tucson, Arizona: derivation, structure, regulation. Condor 76: 184-197. Forbush, E. H. and J. B. May. 1955. A natural history of American birds of eastern and central North America. Revised edition. Bramhall House, New York. 552 pp. Godfrey, W.E. 1966. The birds of Canada. National Museum of Canada Bulletin 203. 428 pp. Haukioja, E. 1969. Weights of reed buntings (Emberiza schoeniclus) during summer. Ornis Fennica 46: 13-21. King, J. K. and D.S. Farner. 1966. The adaptive role of winter fattening in the white-crowned sparrow with comments on its regulation. American Naturalist 100: 403-418. Mcellwraith, T. 1894. Birds of Ontario. William Briggs, Toronto. 426 pp. Middleton, A. L. A. 1977. The molt of the American Gold- finch. Condor. (/n press.) Newton, I. 1967. Adaptive radiation and feeding ecology of some British finches. Ibis 109: 33-98. Newton, I. 1969. Winter fattening in the bullfinch. Physio- logical Zoology 42: 96-107. Newton, I. 1972. Finches. Collins, London. 288 pp. Payne, R. B. 1972. Mechanisms and control of moult. /n Avian biology. Volume II. Chapter 3. Edited by D.S. Farner and J. R. King. Academic Press, New York and London. Raynor, G.S. 1975. Techniques for evaluating and analyz- ing Christmas bird count data. American Birds 29: 626-633. Snyder, L.L. 1951. Ontario birds. Company Limited, Toronto. 248 pp. Snyder, L. L. 1957. Changes in the avifauna of Ontario. /n Changes in the fauna of Ontario. Edited by F.A. Clarke Irwin and 172 Urquhart. Royal Ontario Museum, University of Toronto Press. pp. 26-42. Tast, J. 1968. Changes in the distribution, habitat require- ments and nest sites of the linnet, Carduelis cannabina (L) in Finland. Annales Zooligici Fennici 5: 159-178. Thomas, M. K. 1957. Changes in the climate of Ontario. Jn Changes in the fauna of Ontario. Edited by F.A. Urquhart. Royal Ontario Museum, University of Toronto Press. pp. 59-75. Thomas, M. K. 1968. Some notes on the climatic history of the Great Lakes region. Proceedings of Entomological Society of Ontario 99: 21-31. Thomas, M.K. 1975. Recent climatic fluctuations in Canada. Environment Canada, Climatological Studies Number 28. 92 pp. Tyler, W. M. 1968. Spinus tristis tristis (Linnaeus). Eastern American Goldfinch. /n Life histories of North American THE CANADIAN FIELD-NATURALIST Vol. 91 cardinals, grosbeaks, buntings, towhees, finches, sparrows and allies. Volume |. Edited by O.L. Austin, Jr. Smithsonian Institute Press, Washington, D.C. pp. 447-466. von Haartman, L. 1973. Changes in the breeding bird fauna of northern Europe. /n Breeding biology of birds. Edited by D. S. Farner. National Academy of Science, Washing- ton, D.C. pp. 448-481. Wallace, C. J. 1970. Birds in transition. Massachusetts Audubon 54: 4-10. Ward, P. 1968. Origin of the avifauna of urban and suburban Singapore. Ibis 110: 239-255. Wiseman, A. J. 1975. Changes in body weight of American goldfinches. Wilson Bulletin 87: 390-411. Received 8 September 1976 Accepted 11 February 1977 Notes Incidence of the Dark Color Phase in Tundra and Taiga Populations of Northern Red-backed Voles, Clethrionomys rutilus ARTHUR M. MARTELL Department of Zoology, University of Alberta, Edmonton, Alberta Present address: Canadian Wildlife Service, Environment Canada, c/o Great Lakes Forest Research Centre, Box 490, Sault Ste. Marie, Ontario P6A 5M7 Zimmerman (1961) reported two groups of darken- ing-genes in the genus Clethrionomys: 1, the reces- sive alleles of the agouti series, a (entirely black) and a/t (black with light belly); and 2, the dominant darkening-gene proteus. It is the latter group (pro- teus) that has usually been referred to in the literature as ‘dark phase.’ Dark-phase individuals, those with a distinctly sepia, fuscous, or similarly colored dorsal stripe, have been reported in a number of populations of Clethrionomys gapperi, the southern red-backed vole, but reports of the dark phase in C. rutilus, the northern red-backed vole, are few. Dice (1921) captured two in interior Alaska, and Manning (1956) in his monograph on C. rutilus in Canada reported that only 3 of 386 skins examined were dark phase. Zimmerman (1961) captured a female C. rutilus in Manchuria which produced dark-phase young, but noted that not a single dark-phase individual was found among more than 3000 wild-caught palearctic specimens. Zimmerman’s subsequent breeding experiments with C. rutilus demonstrated some important char- acteristics of the gene proteus. Young proteus have a blackish-brown dorsal stripe when they are in the nest and are in their first subadult coat. At every further molt, however, the amount of black pigment de- creases until, in the last stage, a light sepia-brown color is attained. Individuals in later molts had not previously been recognized as dark-phase. Zimmer- man found that the direction of color change was always from dark to light but that the rate was extremely variable between individuals; some reached the lightest colors within 3-8 months while others were still very dark at 15-16 months of age. Zimmerman suggested that the variability was due to a more rapid fading process in heterozygotes than in homozygotes. Of 83 known proteus examined by Zimmerman only 25% fell into his two darkest color grades, which are likely the only grades previously recognized as dark- phase. The proportion of proteus in natural popu- lations, therefore, has been greatly underestimated. I examined 2607 specimens of C. rutilus captured from tundra and taiga east of the Mackenzie Delta, Northwest Territories from May through October, 1971 to 1973. Taiga collections were made within 6 km southeast of Inuvik (68°00’ N, 133°34’ W) and tundra collections were made at several locations between 68°50’ and 69°40’N, and 133°30’ and 135°10’ W. Neither tundra nor taiga populations offered impressive evidence of being cyclic (Martell 1975). Taiga population indices at the beginning of summer decreased from 1971 to 1973 but late summer indices were higher in 1971 and 1973 than in 1972. Tundra population indices were lower than those in the taiga and were probably similar, and very low, each year at the beginning of summer and were likely similar in 1971 and 1972 but lower in 1973 at the end of summer. Each vole was classed as overwintered (at least 8 months old), early cohort (born in June), or late cohort (born in July and August) based on M2 root development (Martell 1975). Because I was not aware of Zimmerman’s paper at the time I was examining the specimens, only individuals with a distinctly dark dorsal stripe were identified as dark-phase. Those individuals would probably fall into Zimmerman’s two darkest color grades for proteus. Data from taiga collections were examined by month and year for each age class but no significant differences in incidence of the dark phase were found either between years ina given month or between months either within a year or for all years grouped. Therefore the data are presented grouped by year (Table 1). The G-test was used throughout the statistical analysis. Representa- tive specimens have been placed in the Museum of Zoology of the University of Alberta. In the taiga sample there was no significant difference between years in the proportion of dark- 173 174 THE CANADIAN FIELD-NATURALIST Vol. 91 TABLE 1—Percentage of dark-phase Clethrionomys rutilus in collections from tundra and taiga east of the Mackenzie River Delta, Northwest Territories. Sample sizes are in parentheses Location, date Over wintered Taiga May-October 1971 0.0 (147) 1972 0.4 (225) 1973 0.8 (128) 1971-1973 0.4 (500) Tundra June—October 1971-1973 0.0 ( 97) Early-cohort Late-cohort All ages 0.0 (123) 1.6 (317) 0.9 ( 587) 2.9 (137) 2.2 (229) 1.7 ( 591) 6.9 ( 87) 2.1 (243) 2.6 ( 458) 2.9 (347) 1.9 (789) 1.6 (1636) 0.0 (340) 0.0 (534) 0.0 ( 971) phase individuals in the population (0.1 > P > 0.05), suggesting that the incidence of the dark phase is not influenced by population density. Overwintered voles, however, showed a significantly lower frequency of the dark phase than did young-of-the-year (over- wintered vs. early cohort 0.01 > P> 0.001; over- wintered vs. late cohort, 0.05 > P>0.01). Over- wintered animals examined were 8-16 months old and probably contained a higher proportion of faded individuals, which were not recorded, than young-of- the-year which were less than 5 months old. Manning (1956) reported that only 0.8% of the skins he examined were dark-phase. Considering only Manning’s collections from the Mackenzie Delta area, however, dark-phase voles comprised 1.9% of his 107 taiga specimens, not significantly different (0.9 > P > 0.5) from the 1.6% found in this study, and none of his tundra specimens were dark-phase. There was also a total absence of the dark phase in my tundra sample. The difference in incidence of the dark phase between tundra and taiga is significant (P< 0.001). That suggests that selection against the dark phase (at least proteus homozygotes) was greater on the tundra than in the taiga. Peterson (1966) noted that the dark phase occurs throughout the range of C. gapperi but “is apparently clinal, becoming increasingly common in the more northern latitudes.” Because of this trend and knowl- edge of the relation between humidity and darkening factors in other mammals, Zimmerman (1961) sug- gested a positive selective value of the proteus gene in areas with increased humidity. But the incidence of the dark phase in C. rutilus, the more northerly species in North America, appears much lower than that for northern populations of C. gapperi (cf., Harper 1956, 1961; Preble 1908; Smith and Foster 1957). Also the dark phase is extremely rare or absent in at least some tundra populations of C. rutilus. That suggests either that selective factors other than humidity are oper- ating on proteus or that the selective factors are different for C. rutilus than for C. gapperi. One pregnant, apparently red-phase, overwintered C. rutilus, live-trapped near Inuvik, gave birth in the laboratory to three red-phase (1d, 229) and two dark-phase (1d, 12) young. One dark-phase young died at 19 days of age and the second at 42 days of age, but the three red-phase individuals were still healthy more than | year later. The three red-phase young were cross-mated but none of the 18 Fl young produced were dark-phase. The F1 young were cross- mated and were mated with their parents but none of the 28 young produced were dark-phase. That indicates that although the original female captured was a heterozygous proteus, none of the red-phase offspring contained the proteus gene. Those obser- vations are in accord with Zimmerman’s (1961) findings that both homozygous and heterozygous proteus have a dark dorsal stripe at birth and in their first subadult coat. In addition to the proteus individuals, two ap- parently heterozygous agouti late-cohort C. rutilus were taken near Inuvik in January 1972 and August 1972. (The former was taken in additional winter collections by the author.) One tundra population (Tununuk Point: 69°00’ N, 134°40’ W) contained a new color phase. Individuals were of normal color except for the lower back and rump where the tips of many of the guard hairs were sepia. The coloring was similar to the heterozygous agouti individuals but was much less intense. The tundra population was trapped in September and October 1972 and in September 1973, and dark-rump voles were taken in all three samples. In total, 20 overwintered, 95 early-cohort, and 267 late-cohort individuals were examined. None of the overwintered or early-cohort animals were dark-rumped, but six of the late-cohort animals were dark-rumped and two others had dark guard hairs on the upper back and sides as well. Those latter two 1977 individuals still had much less intense coloring than agouti animals. The dark-rump color phase may be similar to the “blackish” color phase found by Harper (1961) in C. gapperi in central Ungava Peninsula. The dark phase (proteus) in Clethrionomys de- serves further attention as a genetic marker because of its obvious phenotype. Such a natural marker could be used to measure changes in the genetic composi- tion of populations between years, particularly in northern C. gapperi populations where 25% or more of the animals may be proteus. I am indebted to T. H. Manning for bringing Zimmerman’s paper to my attention and for his comments on the manuscript. Literature Cited Dice, L. R. 1921. Notes on the mammals of interior Alaska. Journal of Mammalogy 2: 20-28. Harper, F. 1956. The mammals of Keewatin. University of Kansas, Museum of Natural History, Miscellaneous Publication Number 12. 94 pp. Harper, F. 1961. Land and fresh-water mammals of the NOTES 175 Ungava Peninsula. University of Kansas, Museum of Natural History, Miscellaneous Publication Number 27. 178 pp. Manning, T.H. 1956. The northern red-backed mouse, Clethrionomys rutilus (Pallas), in Canada. National Museum of Canada Bulletin Number 144. 67 pp. Martell, A. M. 1975. Demography of tundra and taiga populations of Clethrionomys rutilus. Ph.D. thesis, Uni- versity of Alberta, Edmonton, Alberta. 170 pp. Peterson, R. L. 1966. The mammals of eastern Canada. Oxford University Press, Toronto. 465 pp. Preble, E. A. 1908. A biological investigation of the Atha- basca—Mackenzie Region. North American Fauna Num- ber 27. 574 pp. Smith, D. A. and J. B. Foster. 1957. Notes on the small mammals of Churchill, Manitoba. Journal of Mam- malogy 38: 98-115. Zimmerman, K. 1961. “Proteus,” a new colour gene in bank voles Clethrionomys Tilesius (Mammalia:Roden- tia). Bulletin of the Research Council of Israel 10B: 7-11. Received 11 August 1976 Accepted 9 November 1976 First Record of Atka Mackerel, Pleurogrammus monopterygius (Hexagrammidae), in British Columbia ALEX E. PEDEN British Columbia Provincial Museum, Victoria, British Columbia V8W IAI Pleurogrammus monopterygius (Pallas) is a mid- water greenling (Hexagrammidae) of commercial and sporting value in Soviet (Ruttenberg 1954) and Alaskan waters (Evermann and Goldsborough 1907; Scheffer 1959). Although usually called Atka mack- erel (Miller and Lea 1972, p. 116; Quast and Hall 1972, p. 19), a name recognized in the American Fisheries Society’s, A List of Common and Scientific Names of Fishes (Bailey et al. 1970, p. 57), the name forktail greenling appears in recent literature (Fitch and Lavenberg 1973, p. 131). Previous known distribu- tion for the species is from southeastern Alaska to the Yellow Sea and Sea of Japan (Quast and Hall 1972) and Monterey, California (Miller and Lea 1972). Although its occurrence in British Columbia was expected, there are no documented records. Even though abundant in parts of its geographic range, adequate numbers of specimens for meristic studies are lacking in North American museums (Quast 1964). On 24 August 1976 a specimen 126 mm in standard length was captured after an overnight set of experimental gill nets at the south end of Hunger Harbour, Tasu Sound, Queen Charlotte Islands (52°45’/12” N, 132°01'23” W) by staff of the British Columbia Provincial Museum. The nets were 1.8 m(6 ft) in height and were fished on the bottom between 12.2 m (40 ft) and 36.6 m (120 ft) below 0 tide level. Identification was based on its unnotched dorsal fin, forked tail, and five separate lateral lines. There are 20 spinous dorsal, 26 soft dorsal, 24 anal, and 25 pectoral fin rays on the specimen. The second lateral line had 149 pores and another 16 on the caudal fin. The first, second, and fifth lateral lines terminated posterior to the caudal peduncle. The third lateral line terminated above the 18th anal ray and the fourth ended opposite the tip of the depressed pelvic rays; however, there was a tendency for the fourth to merge with the fifth lateral line on the left side rather than overlap the third as illustrated by Ruttenberg (1955). This specimen 176 THE CANADIAN FIELD-NATURALIST Vol. 91 FiGurE |. The first specimen of Pleurogrammus monopterygius (BCPM 976-1389) from Canadian waters. (British Columbia Provincial Museum catalogue number BCPM 976-1389) is illustrated in Figure | and provides the first published record of Atka mackerel for British Columbia waters. Literature Cited Bailey, R. M., J. E. Fitch, E. S. Herald, E. A. Lachner, C. C. Lindsey, C. R. Robbins, and W. B. Scott. 1970. A list of common and scientific names of fishes from the United States and Canada. American Fisheries Society Special Publication 6. 149 pp. Evermann, B.W. and E.L. Goldsborough. 1907. The fishes of Alaska. Bulletin of the United States Bureau of Fisheries 26: 291-360. Fitch, J. E.and R. J. Lavenberg. 1973. Tidepool and near- shore fishes of California. University of California Press, Berkeley. 156 pp. Miller, D. J. and K.N. Lea. 1972. Guide to the coastal marine fishes of California. Bulletin of the Department of California Fish and Game 157: 1-235. Quast, J.C. 1964. Meristic variation in hexagrammid fishes. Fisheries Bulletin 63(3): 589-609. Quast, J. C. and E. L. Hall. 1972. List of fishes of Alaska and adjacent waters with a guide to some of their literature. National Oceanic and Atmospheric Admin- istration Technical Report, National Marine Fisheries Service Special Scientific Report, Fisheries Series 658: 1-47. Ruttenberg, E.P. 1954. Classification of fishes of the Terpug family (Hexagrammidae). Translated by Lisa Lance and Norman J. Wilimovsky, Fisheries Research Laboratory, Juneau, Alaska, 1958. (Sistema ryb semeys- tua Terpugoykh (Hexagrammidae)). Voprosy Ikhtio- logii 2: 151-155. Ruttenberg, E. P. 1955. On the systematic position of the Terpug genus Pleurogrammus Gill (Pisces, Hexagram- midae). Translated by Lisa Lance and Norman J. Wilimovsky, Fisheries Research Laboratory, Juneau, Alaska, 1958. (O. sistematicheskom polozhenii Terpugov roda Pleurogrammus Gill (Pisces, Hexagrammidae)). Voprosy Ikhtiologii 4: 10-15. Scheffer, B. V. 1959. Invertebrates and fishes collected in the Aleutians, 1936-38. North American Fauna, United States Department of the Interior 61: 365-406. Received 19 October 1976 Accepted 23 December 1976 Apparent Distraction Display by a Barred Owl DAVID M. BIRD! and JO WRIGHT2 'Macdonald Raptor Research Centre, Macdonald Campus of McGill University, Quebec HOA 1C0 766 Cedar Street, Hudson, Quebec JOP 1HO On 18 June 1976 we were searching a section of mixed hardwood forest in Hudson, Quebec for the nest and young of a pair of Barred Owls (Strix varia). As we approached the nest vicinity, the female owl, identified by her higher-pitched voice as compared to that of the male heard on earlier occasions, made her presence known to us by emitting calls described by Eckert and Karalus (1974) as “hoo — hoooo hoo- WAAHHHHhhh, gradually fading away.” She appeared relatively unafraid and remained perched about 20 m high in a beech tree (Fagus grandifolia) maintaining a close watch on us. The male, however, was not observed on this date. Shy versus aggressive behavior towards intruders in the nest area appears to LOTT. vary with individual birds (Dunstan and Sample 1972; Eckert and Karalus 1974). On four occasions during our half-hour search, the female glided downward, distances ranging from 20 to 60 m, to alternate perches about 3 to6 m high in other trees partially hidden from our view by foliage. Immediately upon landing on the perch either facing us or facing away, she spread and quivered both wings and simultaneously uttered a series of chitters and squeals resembling those generally made by begging young. This behavior lasted only 5 to 10 s, after which she crouched on the branch with her wings kept partially opened. When we approached within 20 m of her, the owl returned to one of several high perches located in a central area of roughly 2500 m2. Although we are both experienced field observers, we were fooled on the first two occasions into thinking she had led us to a nest of young or to one of her fledglings begging for food. An intensive search of the area surrounding these perches revealed neither of these and thereafter we restricted our search to the central area. The following day three young Barred Owls were discovered perched high in the trees in the same area occupied by the adult female the previous day. Injury-feigning displays designed to draw intruders away from young have been cited by Bent (1938) for NOTES Wd Great Horned Owls ( Bubo virginianus), Snowy Owls (Nyctea scandiaca), Long-eared Owls (Asio otus), and Short-eared Owls (Asio flammeus), but these displays have never been reported for Barred Owls (M. Fuller, T. Dunstan, personal communication). Of further interest here, however, is that the distraction display was not of an injury-feigning nature, but appeared to simulate the feeding of a young bird by the parent. One other distraction display is reported by Eckert and Karalus (1974) for the Long-eared Owl, that is that the parent attempts to draw intruders away from its young by noisily pretending to catch and kill “some kind of bird as prey.” Only additional observation on distraction displays on any owl species can serve to clarify these behavioral phenomena. Literature Cited Bent, A. C. 1938. Life histories of North American birds of prey. Part 2. Dover Publications, Inc., New York. 482 pp. Dunstan, T. and S. Sample 1972. Biology of Barred Owls in Minnesota. Loon 44: 111-115. Eckert, A. W. and K. E. Karalus. 1974. The owls of North America. Doubleday and Co., Inc., New York. 278 pp. Received 5 July 1976 Accepted 20 December 1976 Natural History of Rock Voles (Microtus chrotorrhinus) in Minnesota ROBERT M. TIMM,! LAWRENCE R. HEANEY,? and DONNA DAY BAIRD Bell Museum of Natural History, University of Minnesota, Minneapolis, Minnesota 55455 'Present address: Department of Entomology, Fisheries, and Wildlife, University of Minnesota, St. Paul, Minnesota 55108 2Present address: Museum of Natural History, The University of Kansas, Lawrence, Kansas 66045 Abstract. A population of rock voles, Microtus chrotorrhinus, which inhabited a large bed of boulders in northeastern Minnesota, was studied in August 1975. The voles did not occupy the entire boulder field, but rather appeared to be restricted to a narrow transition zone between the open rocks and mature forest. An interconnecting system of runways was found in the crevices beneath and between the boulders. Litter size averaged 3.5, with some females producing at least three litters during the breeding season. Females born in late spring produced litters during their first summer. Notes on food habits, activity, parasites, cranial measurements, and associated species are included. The rock vole, Microtus chrotorrhinus, occurs from the Ungava Peninsula to the southern Ap- palachian Mountains, and west along the northern shores of the Great Lakes to Minnesota (Hall and Kelson 1959). Rock voles are restricted to moist rocky habitats in the Canadian and Hudsonian life zones (Kirkland 1977; Linzey and Linzey 1971; Martin 1971a; Doutt et al. 1973; Timm 1974), or more rarely to openings in moist forest (Goodwin 1929; Kirkland 1977). Prior to this study, the rock vole was known in Minnesota from one specimen taken in 1921 near Burntside Lake, St. Louis County (Swanson 1945; Handley 1954), and two taken in 1973 in Cook County (Timm 1974). 178 Study Area and Methods The study area was a long, narrow boulder field, approximately 1.2 km long and 120 m wide, located in sections 19, 20, and 29 of T. 64 N, R. 1 E, Cook County, Minnesota. This open boulder field crosses County Road 12 at 27 km N and 2 km W of Grand Marais, Minnesota, and lies at an elevation of approximately 540 m in a broad valley between two low hills. Frost action associated with retreat of glaciers from this area approximately 9000 years BP (Before Present) is thought to have been responsible for the development of the bed of granophyre and gabbro boulders. Timm (1975) summarized details of climate, vegetation, and mammals in Cook County. The center of the boulder field consisted of exposed rocks occasionally interspersed with small “islands” of shrub vegetation. Dominant vegetation in open rock areas consisted of dry lichens and reindeer moss (Cladonia). The forest surrounding the rock bed was dominated by aspen (Populus tremuloides), paper birch (Betula papyrifera), and black spruce (Picea mariana). Young balsam fir (Abies balsamea) also was common. Thimbleberry (Rubus parviflorus), Clinton’s lily (Clintonia borealis), bunchberry (Cor- nus canadensis), wild lily-of-the-valley (Maianthe- mum canadense), and large-leaved aster (Aster macrophyllus) were the most common herbs. The transition zone between open rocks and forest, ranging from 5 to 10 m in width, was dominated by woody vegetation from | to 3 m in height. Especially common in this zone were alder (Alnus), willow (Salix bebbiana), honeysuckle (Diervilla lonicera), service- berry (Amelanchier), and young balsam fir. Several small plants, especially rose (Rosa acicularis), blue- berry (Vaccinium angustifolium), Clinton’s lily, bunchberry, wild lily-of-the-valley, twin-flower (Lin- naea borealis), and black spruce were common. Boulders covered with moss and leaf litter extended well into dense forest, indicating that the size of the open boulder field has diminished with vegetational succession. Standing water was visible under a small part of the boulder field. Two hundred museum special mouse-traps and 50 Sherman live-traps were baited with peanut butter and oatmeal, and checked four times daily 8-12 August 1975. The 24 rock voles and representatives of all other species trapped were prepared as study specimens and deposited in the Bell Museum of Natural History, University of Minnesota (MMNH). Results and Discussion Habitat Selection All rock voles taken during this study were captured in the boulder field; nearly all were captured in the transition zone between open rocks and mature forest. THE CANADIAN FIELD-NATURALIST Vol. 91 Within this zone most were trapped below the rock surface in cavities between boulders. These cavities were partially filled with soil and were connected to the rock surface and to each other by runways. The southern red-backed vole, Clethrionomys gapperi, was the only other small mammal captured in subsurface runs. The pronounced preference of rock voles for the transition zone probably was due to availability of both preferred food and nesting sites. The subsurface environment presumably assisted in avoidance of both predators and extreme weather conditions. One trap set about 25cm below the surface captured three adult rock voles, indicating multiple use of runways. Reproduction Eleven of 13 adult and subadult females were active reproductively; at least two of these were young-of- the-year as indicated by body size and cranial characters. A mean litter size of 3.5 (N=13) was estimated from counts of embryos, corpora lutea associated with unimplanted embryos, and recent placental scars (see Table 1). These results are similar to data presented by Martin (197la) and Coventry (1937), who found mean litter sizes of 3.7 and 3.6, respectively. Proportionately fewer males than females in our sample were sexually active. Four of the 11 males trapped were in breeding condition as judged by size of the testes and development of the epididymides and seminal vesicles. Average length and width of the testes of these four animals were 12.8 and 7.5 mm, body weight ranged from 32.7 to 43.7 g, and the smallest specimen was 150 mm in total length. The seven non-reproductively active males (testes length X width <5 X 3 mm) varied in weight from 16.2 to 24.8 g. None had a total length greater than 150 mm. Timm (1975) reported that deer mice, Peromyscus maniculatus, southern red-backed voles, and meadow voles, Microtus pennsylvanicus, started breeding in Cook County in May of both 1972 and 1973 and that females of the latter two species may have three litters during the summer. Evidence of three pregnancies (see Table 1) in two large female rock voles in our sample indicated that May or June also may be a typical time for initiation of breeding by rock voles at this locality. Food Habits Most blueberry bushes (both leaves and stems) and Clinton’s lily plants along the narrow margin of the boulder field where rock voles were trapped were heavily browsed by rodents, as indicated by tooth marks. A smaller proportion of wild lily-of-the-valley, bunchberry, and mushrooms appeared to have been 1977 NOTES 179 TABLE 1—Reproductive characteristics of 11 subadult and adult female rock voles from Cook County, Minnesota, collected in August of 1973 and 1975. Code abbreviations are as follows: embs (embryos), CL (corpora lutea associated with unim- planted ova), RS (recent placental scars), OS (old placental scars), L (left uterine horn), R (right uterine horn). Animals are listed in decreasing order of size Catalog Total Weight Number of Litter Comments number length (g) pregnancies size (mm) 12266 160 33.0 2 3RS (ILX2R) Trapped from same locality 40S (2LX2R) in August 1973 12996 158 46.0 3 4CL (1LX3R) 5RS (4LX1R) SOS (2LX3R) 12986 156 40.0 3 CL (unknown #) Well developed mammary 3RS (2LXIR) tissue 6OS (3LX3R) 12985 152 34.9 2 3CL (2LXIR) Lactating 4RS (2LX2R) 12979 152 26.5 2 4 embs (3LX1R) 40S (2LX2R) 12998 147 39.5 2 4 embs (3LXI1R) CL associated with OS (unknown #) embryos = 2LX1R; thus, possibly a case of poly- embryony 12982 147 28.3 D. 3CL (3R) Well developed mammary 30S (2LX1R) tissue 12997 146 36.4 2 3 embs (2LX1R) | resorbing embryo L OS (unknown #) 12978 145 = 2 3 embs (1LX2R) Well developed mammary 5OS (4LX1R) tissue 12983 143 29.6 ! 3RS (2LXIR) Well developed mammary tissue 12981 133 19.1 1 3CL (1LX2R) browsed. Grass was less common and showed little evidence of being grazed. Three rock voles carried plant material in their mouths when captured; one had the partial leaf of a forb, one carried two seeds anda bud, and the third was carrying 3- to S-cm clippings of fresh grass. Captive rock voles consumed blueberry (stems, leaves, and ripe berries), leaves of Clinton’s lily and wild lily-of-the-valley, bunchberry (leaves and Tipe berries), and ripe raspberry (Rubus strigosus), but showed little interest in fresh grasses from the site of capture. A captive subadult male readily consumed all insects presented to him, suggesting that the rock vole may be omnivorous rather than strictly herbiv- orous as previously believed. Activity Patterns Goodwin (1929) and Martin (197la) stated that rock voles are active primarily during daylight hours. This apparently was not the case during our study: of 21 individuals trapped on 9 and 10 August, 7 (33%) were taken between 2300 and 0700 hours (33% of the day); 4 (19%) between 0700 and 1200 hours (21% of the day); 2 (10%) between 1200 and 1800 hours (25% of the day); and 8 (38%) between 1800 and 2300 hours (21% of the day). Timm (1974) reported capturing one rock vole from this population in the early evening and another in the early morning. Thus, it appears that rock voles are active throughout the day and night, but less active during afternoon hours, at least in northern Minnesota during August. Parasites Parasites collected from rock voles during this study include mites, Laelaps kochi and Haemo- gamasus ambulans (Thorell, 1872) [=H. alaskensis Ewing, 1925]; chiggers (Neotrombicula microti and 180 Neotrombicular harperi); ticks (Ixodes angustus); and tapeworms (Cestoda: Hymenolepididae). This record represents the first time Neotrombicula microti has been identified as being parasitic on rock voles. Timm (1974) also reported two species of fleas (Pero- myscopsylla catatina and Megabothris quirini) and one species of mite (Laelaps alaskensis) parasitizing rock voles at this locality. Cranial Measurements Selected cranial measurements (mean and range in millimetres) for four adult females followed by those of four adult males are as follows: greatest length of skull 26.4 (26.2-26.6), 27.1 (26.8-27.4); zygomatic breadth 14.6 (14.4-14.7), 15.1 (14.4-15.6); inter- orbital constriction 3.6 (3.5-3.6), 3.7 (3.5—3.7), length of nasal bones 7.4 (7.2-7.5), 7.6 (7.4-7.9); length of maxillary toothrow 6.2 (5.8-6.4), 6.3 (6.2-6.5). These measurements are larger than corresponding measurements reported by Komarek (1932) for specimens of the same subspecies, M. chrotorrhinus chrotorrhinus, from the eastern part of their range. Associated Species Nine other mammalian species (followed by the number of each trapped) were taken from the study area in 1975: short-tailed shrew, Blarina brevicauda (1); eastern chipmunk, Tamias striatus (1); least chipmunk, Eitamias minimus (7); red squirrel, Tam- lasciurus hudsonicus (1); deer mouse (6); southern ted-backed vole (48); meadow vole (2); southern bog lemming, Synaptomys cooperi (5); and ermine, Mustela erminea (1). Timm (1974) also captured the arctic shrew (Sorex arcticus), masked shrew (S. cinereus), and woodland jumping mouse (Napaeo- zapus insignis) at this site. Several mink, Mustela vison, were sighted in the vicinity in 1973 and 1975. The presence of four species of microtine rodents at this site is of interest in regard to competition and competitive exclusion in small mammals. Martin (1971b) found no reports of the meadow vole in habitat occupied by the rock vole. Our two meadow voles were non-breeding subadult males and were trapped in the open rocks. Breeding populations of meadow voles apparently occur in the area and individuals disperse to the rock outcrops, but meadow voles have not successfully colonized the boulder field. Five southern bog lemmings were trapped in the transition zone and adjacent mature forest. One of these was removed from a trap at which a rock vole and a southern red-backed vole had been captured previously. The presence of a pregnant adult female suggests a resident population of southern bog lemmings at the site, and indicates at least some overlap of habitat use by bog lemmings and rock THE CANADIAN FIELD-NATURALIST Vol. 91 voles. Southern bog lemmings appear to have a broader habitat range, utilizing forest areas as well as the transition zone. The southern red-backed vole was an abundant small mammal both in the preferred habitat of the rock vole and in the adjacent forest. It was the only other microtine taken from subsurface runs, and in at least three instances was taken from traps that also caught rock voles. In addition to being active at the same time, red-backed voles probably eat many of the same foods, and breeding by the two species takes place during the same times of the year. No fighting occurred in laboratory investigations of behavior using a single subadult male rock vole and three adult red-backed voles, but the rock vole appeared to be dominant. Red-backed voles were aggressive when handled however, whereas rock voles were docile. Conclusions The ecology of the rock vole suggests several questions for future investigation. Because all popu- lations of rock voles reported seem to be small and isolated from other populations, it would be espe- cially interesting to examine patterns of genetic variability. What allows rock voles to compete successfully with several other microtine rodents, especially red-backed voles, which appear to occupy a similar niche in this environment? Is their non- aggressive behavior and low litter size a response to a relatively stable and predator-free subterranean living situation? Do they show cyclic patterns of population fluctuation? The rock vole is rare and remains poorly known in Minnesota. As the forest encroaches upon the rock bed, habitat available for rock voles is slowly decreasing; however, a more immediate threat to the population is destruction of habitat for timber harvest. Steps should be taken to insure that this site is protected and that scientific investigations on this population be conducted in such a manner as not to threaten its future. We acknowledge the valuable criticism of E. C. Birney, G. E. Glass, R. S. Hoffmann, R. P. Lampe, and J. A. Thomas on various drafts of the manu- script. We also thank R. C. Bright and R. M. Schaefer for technical assistance. Literature Cited Coventry, A. F. 1937. Notes on the breeding of some ‘Cricetidae in Ontario. Journal of Mammalogy 18: 489-496. Doutt, J. K., C. A. Heppenstall, and J. E. Guilday. 1973. Mammals of Pennsylvania. Pennsylvania Game Com- mission, Harrisburg, Pennsylvania. 288 pp. Goodwin, G. G. 1929. Mammals of the Cascapedia Valley, Quebec. Journal of Mammalogy 10: 239-246. 1977 Hall, E.R. and K.R. Kelson. 1959. The mammals of North America. The Ronald Press Company. Volume 2. pp. vilit547-1083+79. Handley, C. O., Jr. 1954. Phenacomys in Minnesota. Jour- nal of Mammalogy 35: 260. Kirkland, G. L., Jr. 1977. The yellow-nosed vole, Microtus chrotorrhinus (Miller) (Mammalia: Rodentia) in West Virginia. Annals of Carnegie Museum, Pittsburg. (/n press.) Komarek, E. V. 1932. Distribution of Microtus chrotor- rhinus, with description of a new subspecies. Journal of Mammalogy 13: 155-158. Linzey, A. V. and D. W. Linzey. 1971. Mammals of Great Smoky Mountains National Park. University of Ten- nessee Press, Knoxville, Tennessee. 114 pp. Martin, R. L. 197la. The natural history and taxonomy of the rock vole, Microtus chrotorrhinus. Ph.D. thesis, University of Connecticut, Storrs, Connecticut. 164 pp. NOTES 181] Martin, R.L. 1971b. Interspecific associations of rock voles. Beta Kappa Chi Bulletin 30(2): 5-7. Swanson, G. 1945. A systematic catalog of the mammals of Minnesota. In The mammals of Minnesota. Edited by G. Swanson, T. Surber, and T. S. Roberts. Technical Bulletin of the Minnesota Department of Conservation 2: 52-105. Timm, R. M. 1974. Rediscovery of the rock vole (Micro- tus chrotorrhinus) in Minnesota. Canadian Field-Natura- list 88: 82. Timm, R.M. 1975. Distribution, natural history, and parasites of mammals of Cook County, Minnesota. Occasional Papers Bell Museum of Natural History, University of Minnesota 14: 1-56. Received 26 August 1976 Accepted 21 November 1976 Disorientation in Ringed and Bearded Seals THOMAS G. SMITH! and JIMMY MEMOGANA2 'Fisheries and Marine Services, Department of Fisheries and the Environment, Arctic Biological Station, P.O. Box 400, Ste. Anne de Bellevue, Quebec H9X 3L6 2Holman, Northwest Territories The phenomenon of seals lost on land or unable to find access to the water through the frozen ice cover is well documented for some antarctic localities (Stirling and Rudolph 1968; Stirling and Kooyman 1971). Although it is common knowledge among the Inuit that seals get lost, few reports exist of such occurrences for the Arctic. Freuchen and Salomonsen (1958) and Freuchen (1935) note that walruses (Odobenus rosmarus) and ringed seals (Phoca hispida) sometimes get caught out of the water by freezing ice. This note documents several instances where both live or dead ringed seals and bearded seals (Erignathus barbatus) have been found on the land or away from access to water. Data were collected over an 8-year period in the Home Bay region, east Baffin Island, and in the Holman region on the west coast of Victoria Island, Northwest Territories. Ringed Seals On the Sea Ice June 1968. During a hunt of hauled-up seals on the flat ice northeast of Ekalugad fiord (68°40’ N, 65°10’ W) the trail of a lost seal was found. By following the tracks for approximately 1.6 km over the flat sea ice, we located and collected a ringed seal pup (0+ years). 14 June 1972. A track was followed for at least 6.6 km until the ringed seal pup (0+) was found on the flat sea ice near Tluvilik (70°30’ N, 116°30’ W) southeast of Holman, North- west Territories. Late April or early May 1973. Approximately 25 kmto the west of Holman (70°57’ N, 118°25’ W) on the shoreline, an adolescent ringed seal was found. The track of the seal was found on the ice, trailing inland for a short distance and then returning to the sea ice. The land at this location rises as a gentle slope from the ocean. This area of shoreline also shows a consistent opening and closing of tidal cracks during the fast-ice season. The seal had apparently been out of the water for some time, since its hindquarters were frozen. The ventral skin surface was badly worn indicating that the animal had travelled a considerable distance. There was some evidence of bite marks in the axilla and on the hind flippers. On the Land 5 May 1972. A yearling male ringed seal was sighted approximately 18km inland northwest of Holman (70°47’ N, 117°49’ W) near Okotitak Lake. The seal was alive and still moving. It had very badly worn areas of skin ventrally. Another seal was found crossing Irkaharvik Lake (70°52’ N, 117°56’ W) approximately 17 km inland north- east of the village of Holman. The exact year of occurrence is not remembered but the seal was found in late March or early April. Early June 1973. A live adolescent ringed seal was found approximately 150 m inland on the south shore of Prince 182 Albert Sound (70°30’ N, 116°32’ W) near Cape Baring. The condition of the seal indicated it had recently come ashore. The shoreline is very low, and the boundary between sea ice and land, covered by snow, imperceptible to the human eye. 28 August 1973. A whole skeleton of a medium-sized ringed seal was found in a high valley approximately 60 m above sea-level on an island in Prince Albert Sound (70°30’ N, 116°30’ W). The valley could have been reached from the end of a long crooked inlet penetrating into the island for a distance of approximately 2 km. It is unlikely that the carcass would have been left by the Inuit since no one traps inland on these islands. Bearded Seal 27 March 1974. A 0+-year-old bearded seal measuring 100 cm nose to tail, 115 cm maximum girth, was found frozen into the surface ice of a lake. The lake was located at 71°10’ N, 118°00’ W, 52 m above sea-level and connected to Minto Inlet by a small steeply rising creek. In its frozen condition the specimen displayed the “hunched back” posture seen in distressed seals. Its foreflippers were folded ventrally and the hind flippers curled toward each other. The skin on the sternum and the axilla region was worn indicating that the seal had travelled over land or ice. The vibrissae also showed excessive wear. The blubber layer was virtually nonexistent suggesting the seal had not fed well for some time. Examination of the stomach and intestines showed the seal had attempted to nourish itself by eating arctic willow and grasses. A quantity of brown matter, probably earth, was also found. No evidence was seen of fish or invertebrate remains. It is unclear whether the seal had journeyed up to the lake after freeze-up or if it had gone up the river during the open-water season the previous summer. Discussion Instances of ringed seals found away from access to water result from seals moving away from their exit holes or being frozen out of the hole or crack they had emerged from. Both cases of ringed seals found moving over the ice during the period when exit holes no longer freeze quickly involved young-of-the-year (0+ age). Observa- tions of pups still with their mothers during the haul- out period from May through June showed them to move further away from the breathing holes than the adult seal. If visual clues are used to locate the hole it is not surprising that pups frequently become dis- oriented. At this time of the year, however, a large number of breathing holes and cracks are present so that many of the disoriented pups probably regain access to the water. The air temperatures are also high enough so as not to cause many deaths through freezing. A different situation exists with adolescent seals (1+ to 6+ years old) found relatively frequently on the ice or the land during the early spring months of March and April. Here loss of access to the water usually results in death. The reasons for the initial emergence THE CANADIAN FIELD-NATURALIST Vol. 91 onto the ice, and why the seal becomes frozen out are not at all clear. The presence of bite marks and the often poor nutritional state of the seals may suggest some form of intraspecific competition being in- volved. Whether this is connected with territoriality or simple competition for food is not known. It is also possible that diseased and sick seals emerge from the water simply to rest and because of their weakened state get frozen out. We have also found several dead adolescent seals inside haul-out lairs under the snow. The actual movement of seals from the water or sea ice onto the land may be explained in several ways. During the snow-covered period it is very difficult to distinguish visually between the sea ice and the land along low-lying coastlines. If in fact seals are using visual clues as they are trying to relocate a hole or crack, they could easily move onto the land without knowing it. In the summer open-water period, ringed seals and bearded seals occasionally move into small rivers. In one instance a ringed seal pup (0+ age) was seen swimming up the small Anialik River (70°34’N, 116°57’ W). The pup continued to swim into the current even after it had seen me wading out to it from a point upstream. It remained undaunted in its attempt to swim up the river against the strong current until I captured it. Such orientation into the currents of small rivers and movements up these streams might lead to the occasional disorientation and emergence onto the land in ringed seals. Bearded seals in the summer months are known occasionally to haul out onto the land to bask. This habit might also occasionally lead to disorientation inland among the younger inexperienced seals. Acknowledgments Thanks are due to my friends the hunters and trappers of Holman who keep me informed about the wildlife of the area. Mr. Brian Glandfield, formerly of Holman, provided useful information. Literature Cited Freuchen, P. 1935. Mammals: Part II. Field notes and biological observations, /n Report of the fifth Thule expedition 1921-24 Volume II. Pp. 68-278. Freuchen, P. and F. Salomonsen. 1958. The Arctic year. G. P. Putnam’s Sons, New York. 438 pp. Stirling, I. and E. D. Rudolph. 1968. Inland record of a live crabeater seal in Antarctica. Journal of Mammalogy 49: 161-162. Stirling, I. and G. L. Kooyman. 1971. The crabeater seal (Lobodon carcinophagus) in McMurdo Sound, Ant- arctica, and the origin of mummified seals. Journal of Mammalogy 52: 175-180. Received 26 November 1976 Accepted 7 January 1977 1977 NOTES 183 Breeding Status of the Say’s Phoebe in Manitoba RICHARD W. KNAPTON Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Godfrey (1966) includes southwestern Manitoba in the breeding range of the Say’s Phoebe (Sayornis saya) on the basis of Taverner’s (1927) report of the species nesting several times in the neighborhood of Aweme, which was formerly near Treesbank, as reported to Taverner by Norman Criddle, then of Aweme. There is, however, no reference on file at the Manitoba Museum of Man and Nature, Winnipeg, of the species having nested at Aweme, and moreover the Manitoba Museum Field Check-list of Manitoba Birds (1974) classes the species as of irregular occurrence in Manitoba. Further, Brazier (1956) could find no published record of the _ bird’s occurrence in Saskatchewan farther east than Regina, although it was recorded in open country near Moose Mountain, Saskatchewan in 1959 (Nero and Lein 1971). In this paper, I show that the Say’s Phoebe has nested several times in recent years in southwestern Manitoba, and that it is a rare but regular breeding species in the province. Prior to the 1970s, the only nesting record in Manitoba, apart from those reported by Taverner (1927), was in 1946 at Oak Lake where Herman Battersby (personal communication) located a pair nesting in an old house, the pair raising five young. In 1972 a pair repeatedly tried to nest at the Customs Office on Highway 256, 11 km southwest of Lyleton. The pair finally constructed a nest on the third attempt, but on 19 June the female was found dead beneath the nest, which contained three eggs. The bird is now specimen 3229 in the Manitoba Museum of Man and Nature. In both 1973 and 1974, a pair returned to the same location for the summer, but no evidence of nesting was found, although a family party of five birds on 16 July 1974 indicated a successful nesting somewhere in the immediate area (J. L. Murray, personal communication). On 26 May 1974, I located a pair of Say’s Phoebes at an abandoned farmhouse 7 km southeast of Lyleton, and on 2 June R. F. Koes and I found the nest being built inside a nearby barn. The nest contained five eggs on 21 June, and the pair succeeded in raising at least three young. The nest is now in the Manitoba Museum of Man and Nature, catalogue number 1|.21-338. In 1975 and 1976, Say’s Phoebes returned to the farmhouse. No nest was located in 1975, but in 1976.a nest was found on 7 June in an old Barn Swallow (Hirundo rustica) nest, containing four young phoebes, three of which were subsequently reared (personal observations). Thus, in five consecutive years, Say’s Phoebes have bred or have been present through the summer in the vicinity of Lyleton, in the extreme southwestern corner of Manitoba. This part of the province is mixed-grass prairie, arid and hot in summer, with numerous abandoned farms, 1.e., one type of habitat Say’s Phoebes will inhabit (Bent 1963). Away from this habitat in Manitoba, the Say’s Phoebe is very rare: indeed, there are no sight records north of Brandon (Lawrence 1934) and Oak Lake (D. R. M. Hatch, personal communication), nor east of Aweme (Criddle 1913). In adjacent counties in the North- western Drift Plain in North Dakota, the species is classed as uncommon, although several nests with dependent young have been located in the last 20 years (Stewart 1975). The Say’s Phoebe then is a rare but regular breeder in Manitoba, with individual pairs widely dispersed and with probably no more than 10 pairs per year. I thank W. E. Godfrey and H. W. R. Copland for assistance in locating past records, and several people who supplied me with information from their own personal records, in particular H. Battersby, D. R. M. Hatch, and J. L. Murray. Literature Cited Bent, A.C. 1963. Life histories of North American fly- catchers, larks, swallows, and their allies. United States National Museum Bulletin 179. 555 pp. Brazier, F. H. 1956. Say’s Phoebe in Saskatchewan. Blue Jay 14: 91-92. Criddle, N. 1913. New or rare bird records from Manitoba, 1912. Ottawa Naturalist 26: 126-127. Godfrey, W.E. 1966. The birds of Canada. National Museum of Canada Bulletin Number 203. 428 pp. Lawrence, A. G. 1934. Chickadee notes 688. Winnipeg Free Press, | June 1934, Winnipeg. Nero, R.W. and M.R. Lein. 1971. Birds of Moose Mountain, Saskatchewan. Saskatchewan Natural History Society Special Publication 7. Stewart, R.E. 1975. Breeding birds of North Dakota. North Dakota State University, Fargo, North Dakota. Taverner, P. A. 1927. Some recent Canadian records. Auk 44: 217-228. Received 24 October 1976 Accepted 17 January 1977 184 THE CANADIAN FIELD-NATURALIST Vol. 91 Le Grand Cormoran (Phalacrocorax carbo) en Hiver, le Long des Cétes de la Péninsule de Gaspésie, Québec GILLES CHAPDELAINE Service canadien de la faune, Région du Québec, Edifice “A”, 4e étage, 2700 boul. Laurier, C.P. 10100, Ste-Foy, Québec GIV 4H5 Depuis 1974, des recensements annuels réalisés par le Service canadien de la faune le long des cétes de la péninsule de Gaspésie permettent de mieux connaitre la distribution hivernale de plusieurs espéces d’oiseaux aquatiques. La présence du Grand Cor- moran (Phalacrocorax carbo), parmi les espéces inventoriées, ne manque pas d’intérét puisque sa distribution en hiver est peu documentée et fut considérée jusqu’a ce jour comme inusitée au Parc national de Forillon (Ouellet 1975), bien que Palmer (1962, p. 321) Pait déja inclus comme résident d’hiver en Gaspésie. Erskine (1972) a déja revisé l’aire dhivernage de cette espéce qui se situe principale- ment le long des cétes du Maine et du Massachusetts ainsi qu’en quelques concentrations mineures en Nouvelle-Ecosse (Ross 1974) et au sud-ouest du Nouveau-Brunswick. Les données consignées ici (Tableau 1) proviennent d’inventaires réalisés durant la derniére semaine de janvier en 1974 et la premiere semaine de février en 1975 et 1976. Les conditions climatiques difficiles dans lesquelles les observations de 1974 ont été effectuées, ne nous ont pas permis de distinguer avec satisfaction s'il s’agissait de Phalacrocorax carbo ou TABLEAU 1—Dénombrement du Grand Cormoran le long des cétes de la péninsule de Gaspésie en hiver en 1974, 1975 et 1976 Lieu 1974 1975 1976 Maria 0 16 46 Cap noir a Bonaventure 28 0 0 Hope a Shigawake I) 26 1 Port-Daniel 2 0 44 Newport a Pabos 0 Grande-Riviére a Cap d’Espoir 1 27 Total 56* 72 102 *Lidentification du Grand Cormoran n’a pu étre certifiée en 1974 en raison des conditions de température inclémentes, mais il s’agissait probablement de cette espéce. P. auritus bien que la présence de ce dernier elt été assez inusitée. De meilleures conditions d’observation en 1975 et 1976 ont permis d’identifier tous les individus comme étant des Grands Cormorans. I] est donc probable que les individus observés en 1974 appartenaient aussi a l’espéce P. carbo. Le nombre maximum de 102 Grands Cormorans obtenu en 1976 est inférieur aux estimés que Ross (1974) a rapportés sur la céte de la Nouvelle-Ecosse, soit 390 par un inventaire au sol et 674 par un inventaire aérien. Cependant, nous croyons que le nombre maximum qui apparait ici pourrait étre supérieur si un inventaire aérien était réalisé. Ainsi, a quelques reprises on a vu des individus surgir du rebord de la falaise comme s'il s’agissait d’oiseaux qui quittent un reposoir. Ces endroits ainsi que plusieurs autres sites disponibles pour le Grand Cormoran sont impossibles a inventorier au sol. En 1976, on'a remarqué deux groupes importants qui se reposaient sur le bord de la glace, soit 46 a cap Maria et 44 a Port-Daniel. Cette fagon typique de se rassembler a aussi été observée en 1974 et 1975 chez des groupes plus restreints en divers endroits de la céte. Il est a remarquer qu’aucune observation de cette espéce ne fut consignée entre Percé et Cap-des-Rosiers au cours de ces trois inventaires annuels, comme onl’a noté précédemment (Ouellet 1975). Je remercie Austin Reed et André Bourget d’avoir porté attention a la présente communication. Références Erskine, A.J. 1972. The Great Cormorants of eastern Canada. Canadian Wildlife Service Occasional Paper Number 14. 21 pp. Ouellet, H. 1975. Contribution a l’étude des oiseaux d’hiver au Pare national de Forillon, Québec. Revue Géo- graphique de Montréal 29(4): 289-304. Palmer, R.S. (Rédacteur). 1962. Handbook of North American birds. Volume |, Loons through Flamingos. Yale University Press, New Haven and London. 567 pp. Ross, R. L. 1974. Notes on wintering Great Cormorants in Nova Scotia. Canadian Field-Naturalist 88(4): 493- 494. Recu 6 Juillet 1976 Accepté 21 Novembre 1976 1977 NOTES 185 Records of the Boreal Toad from the Yukon and Northern British Columbia FRANCIS R. COOK Herpetology Section, National Museum of Natural Sciences, Ottawa, Ontario Logier and Toner (1961, pp. 27-28) list and map Prince William Sound and Yujutat in Alaska and Telegraph Creek, Lake Tetana, Hudson Hope, and Tupper Creek in British Columbia as the northern boundary of the range for the Boreal Toad, Bufo boreas boreas. Stebbins (1966, Map 31) apparently followed these points in drawing his northern range limit for the subspecies. Herreid (1963) reported specimens from the Liard Hot Springs about 400 km (250 mi) northeast of the range shown by the above authors. Unnoted by these authors, however, was a specimen collected between | and 10 July 1948, by William Mason at Whitehorse in the Yukon Territory and deposited in the American Museum of Natural History collection (AMNH 54146, 40.3 mm snout- vent length). This is the first record of the species from the Yukon Territory. Several additional unreported northern records are catalogued in the Herpetology Section collections of KIA 0M8 the National Museum of Natural Sciences. A speci- men (NMNS 5864, 41.5 mm snout-—vent length) from North Toobally Lake (60°21’ N, 126° 15’ W, elevation 660 m (2200 ft) collected by P. M. Youngman and G. Tessier on 16 July 1961 is the only other record of the species from the Yukon Territory. It extends the known range some 80 km (50 mi) north of the Liard locality. The field notes of these collectors indicate that the specimen was taken at their campsite on the east shore of the lake, near its south end. This is only the second amphibian species definitely known to occur in the Yukon. The Wood Frog, Rana sylvatica, has been taken at many localities in the territory (Logier and Toner 1961; NMNS collections). One other anuran, the Western Spotted Frog, Rana pretiosa, is reported almost at the Yukon border at Lake Bennett (Carl 1943, p. 50) in northwestern British Columbia. A Boreal Toad (NMNS 2170, 71.0 mm) from Mile 46 of the Haines Road (north of Haines, Alaska but in British Columbia) was collected i; y, CY 4 Z, j x by v¢ GU WSU BAY FIGURE 1. Map of known range of the Boreal Toad, Bufo boreas boreas, in northern British Columbia, Yukon, and Alaska. The hatching represents the range as depicted by Stebbins (1966); open circles are records shown in Logier and Toner (1961); partly open circles are records given by Herreid (1963) and Cowan (1936); solid circles are new records. 186 by W. E. Godfrey on 30 July 1949 and is the first record of this species for this northwestern projection of British Columbia into Alaska. Deirdre Griffiths (personal communication, 1973) has provided a sight observation of another at Mosquito Lake Camp- ground, 3.2 km (2 mi) east of Mile 27.2 on the Haines Highway, on 12 July 1972. Overlooked by Logier and Toner (1961) is a record, given by Cowan (1936, p. K19), from Atlin near the Yukon border in western British Columbia. Hugh S. Bostock (personal communication to J. S. Bleakney, NMNS files, 1952; and to FRC, 1972) reported that toad eggs were observed to be abundant in pools on the south side of Willison Bay (134°10’ N, 59°15’ W), Atlin Lake 6 to 8 July 1952. Numerous tadpoles were seen in these ponds at this locality during this time. He was informed that toads also inhabit the area around the warm spring (at Warm Bay) (133°30’ N, 59°22’ W) about 20 km (12 mi) by road south of the town of Atlin, on the east side of Atlin Lake. Ben-My-Cree (134°29’ N, 59°19’ W) on Tagish Lake, a few miles northwest of Willison Bay is a similar indentation into the Coast Mountains and would seem likely to be in the range of these toads as it too is a sheltered area where snow comes early in the fall and accumulates to depths of 6m (20 ft) and more. In view of these records Bufo boreas should be looked for in the western portion of the Yukonas well. Three additional unreported collections are from the Liard Hot Springs area. NMNS 2070 (four speci- mens: 69, 68.5, 67, 52.5 mm) was taken at “Liard River and Alaska Highway, mi. 213 N. of [Fort] Nelson, B.C.” by A. L. Rand on 13 August 1943. NMNS 2077 (two specimens: 21.0, 16.5 mm) was taken at “Mile 213, Tropical Valley, Alaska Highway, B.C.” also by A.L. Rand on 2 June 1944. NMNS 6513 (three specimens: 60.5, 58.0, 54.5 mm) was taken at the Liard Hot Springs by S. D. MacDonald on 31 May 1962. These specimens pre- date the three specimens taken 4 September 1962 by Herreid (1963) from this area and substantiate a natural population in the area. One additional collection falls between the Liard locality and the record from the Yukon. NMNS 5863 (67 mm) was taken at “Smith River,” British Columbia, by P. M. Youngman and G. Tessier on 9 July 1961. Field notes indicate these specimens were collected near the Smith River airport. All of these northern records seem associated with valleys which probably accumulate an early deep snow cover which prevents deep frost penetration and assures safe terrestrial hibernation. Bostock (personal communication) has informed me that at Ben-My- Cree on Tagish Lake the snow cover reached 6 m (20 ft) or more and that there appeared to be no permafrost. He has cited an occasion when the owners THE CANADIAN FIELD-NATURALIST Vol. 91 (Mr. and Mrs. Partridge) of the Ben-My-Cree home- stead left their vegetables in their garden over winter in the ground and the next spring found their potatoes and other roots had not frozen at all. Although many records are often from the vicinity of hot springs, Deirdre Griffiths’ (personal communication, 1973) points out that the individual she observed at Mosquito Lake was frequenting a little spring used for the camp water supply. Water from this spring was very cold. It is reasonable to suggest that B. boreas may have continuous populations connected through favorable valleys in the northern portion of the range. It may be unnecessary to postulate as Herreid (1963) did, that the Liard population is considerably separated from its nearest neighbor and ascribe survival solely to conditions induced by the presence of hot springs, as heavy early snow accumulation and lack of perma- frost may be equally important. Acknowledgments My gratitude to the several collectors who have deposited specimens in the Herpetology Section, to C. G. van Zyll de Jong for making the field notes of A.L. Rand, P.M. Youngman, and G. Tessier available to me from the files of the Mammalogy Section, National Museum of Natural Sciences, to Richard G. Zweifel, Department of Herpetology, American Museum of Natural History for loaning the specimen from Whitehorse (examined 17 February 1977), and to Deirdre Griffiths for observations. Special thanks are due to Hugh S. Bostock, formerly of the Geological Survey of Canada, now retired, who contributed stimulating observations and comments and suggested important changes in an earlier version of this manuscript. James A. Johnston, Herpetology Section, NMNS, prepared the map. Literature Cited Carl, G. Clifford. 1943. The amphibians of British Colum- bia. British Columbia Provincial Museum, Handbook Number 2. 62 pp. Cowan, Ian McTaggart. 1936. A review of the reptiles and amphibians of British Columbia. Report of the Provincial Museum for 1936, British Columbia. pp. K16-K25. Herreid, Clyde F., II. 1963. Range extension for Bufo boreas boreas. Herpetologica 19(3): 218. Logier, E. B.S. and G. C. Toner. 1961. Check list of the amphibians and reptiles of Canada and Alaska. Royal Ontario Museum, Life Sciences Division, Contribution 53. 92 pp. Stebbins, Robert C. 1966. A field guide to western reptiles and amphibians. Houghton Mifflin Company, Boston. 279 pp. Received 2 July 1975 Accepted 6 August 1975 Updated and resubmitted 23 February 1977 M7 NOTES 187 Predatory Behavior by Common Grackles A. L. A. MIDDLETON Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1 On 17 July 1975 I received a call from a worried neighbor in Guelph, Ontario, who was concerned over the number of mutilated House Sparrow (Passer domesticus) carcasses she was suddenly finding close to her bird feeders. In most cases the carcasses were beheaded and the skin torn from around the shoulders, but in a few instances the head was still attached to the body, the skull was openand the brain removed. A watch was immediately set at the feeding station, which at this time of year was frequented mainly by House Sparrows, Starlings (Sturnus vulgaris), and the Common Grackle (Quiscalus quiscula). During the first day of observation a male grackle was seen to attack a House Sparrow that was feeding on the ground with some other sparrows. As the other birds flushed, the grackle knocked the victim to the ground, held it there and repeatedly pecked at the sparrow’s head. The sparrow was quickly killed, the cranium opened, and the brains eaten. Thereupon, the grackle paid no further attention to the mutilated carcass and flew off, leaving the body. Following this attack food was withdrawn from the feeding station and no further reports of killings were received. Because of the concern of my neighbor, no attempt was made to trap and mark the grackles involved. Although not verified, it is possible that all the killings were the act of a single bird, since 15 sparrows had been killed in the same fashion during an | 1-day period. Mayfield (1954) observed a grackle killing a House Sparrow in the fashion described above and discovered a headless carcass nearby, and Gross (1958) reports several instances of predatory behavior by individual grackles. Thus the development of predatory behavior by individual Common Grackles, particularly when prey is abundant and can be easily captured, may be more common than suggested by the isolated reports elsewhere in the literature (see Laporte 1974). Literature Cited Gross, H. M. 1958. Bronzed Grackle. Jn Life histories of North American blackbirds, orioles, tanagers and allies. Edited by A.C. Bent. United States National Museum Bulletin. pp. 395-421. Mayfield, H. 1954. Grackle kills English Sparrow. Wilson Bulletin 66: 271. Laporte, P. 1974. Common Grackle kills a Barn Swallow. Wilson Bulletin 86: 477-478. Received 8 September 1976 Accepted 23 December 1976 The Function of the Bark Call of the Red Squirrel GARY F. SEARING LGL Limited, Environmental Research Associates, 10110 - 124 Street, Edmonton, Alberta TSN 1P6 Alarm calls that may warn other members of the same species of the presence of potential predators have been reported for several mammalian species; these include Cynomys ludovicianus (King 1955), Spermophilus paryii (Melchior 1971; Carl 1971), Ochotona princeps (Markham and Whicker 1973), Alces alces (De Vos 1958), and others. An alarm function of such calls is well documented for several species, but is still conjectural for others. Calls of the red squirrel (Tamiasciurus hudsonicus) have been described previously by Klugh (1927), Smith (1963, 1968), Embry (1970), and Nodler (1973); the last two authors have discussed the function of several vocalizations. The commonly heard “scold- ing” or “bark” call of the red squirrel has been classified as an alarm call by Smith (1963) who felt that the call served to warn others of predators. Nodler (1973) proposed a less specific purpose for the call; she stated that it was a call of stress or release from a stressful situation. While I was studying the aggressive behavior and population dynamics of red squirrels in interior 188 Alaska, 10 encounters by red squirrels with avian and mammalian predators were observed. An additional seven encounters have been related to me or reported in the literature. The vocal reactions of squirrels to predators help to clarify the function of the bark call of the red squirrel. Table | presents a summary of the vocal behavior of red squirrels during the 17 predator- prey interactions. In 10 of the 17 encounters (59%), the squirrel’s initial response was to remain quiet. Only after the potential predator left did these squirrels give the bark (9 of 10 instances). In five cases the squirrels gave a bark initially; three of these involved a marten. The remaining two cases involved squirrels attacked away from cover by Goshawks. The reactions of the squirrels attacked by Goshawks were similar, and involved fleeing to cover while giving a “rapid squeak” call (a series of high-pitched, rattled notes approxi- mating a succession of barks). Neighboring squirrels never responded with a bark to the bark of another squirrel that had been attacked by a predator, nor did the movement of a predator elicit the bark from squirrels within its path. It appeared that when a squirrel was seriously threat- ened by a predator it was quiet. After the predator left the bark was commonly used, but did not appear to be a specific call of alarm since the squirrels that would have been warned by such a call had already been exposed to the predator. In order to test the response of red squirrels to the bark, I conducted 14 playback experiments using recorded barks. Squirrels responded with bark and “chir” calls (see Klugh (1927) for a description of the chir call) when the speaker was located within 20 m of the midden (four of five). When the speaker was 20 to TABLE 1—Vocal responses of red squirrels to predators* THE CANADIAN FIELD-NATURALIST Vol. 91 30 m away, squirrels tended to give the rapid squeak and chir calls (four of five). When the speaker was beyond 30 m, squirrels gave no vocal response (four of four). In contrast, chir-call playbacks (Searing 1975) frequently elicited chir calls from squirrels when played more than 30 m from the midden. But the fact that the bark elicited a chir call from squirrels when played within 30 m of a squirrel’s midden leads me to believe that this call may contain some aggressive components. It is my impression that the bark is given more frequently when squirrels are not at peak aggressive levels (i.e., during winter) and during moderately aggressive encounters. It is also significant that the chir, an aggressive call (Searing 1975), is given to intruding people, i.e., potential predators. Here again the squirrel usually remains quiet until after the intruder has left, whereupon it usually gives a chir call or less frequently the bark. The fact that these two calls are often used in similar situations further supports my conclusion of a partially common function. Balph and Balph (1966) reported that the aggressive calls of Uinta ground squirrels (Spermophilus armatus) were also used in response to predators and evoked general alertness by all squirrels. My experiments with playbacks of the bark did not yield alert responses by red squirrels. Nodler (1973) stated that the bark was used by subordinate squirrels during border disputes but never by the dominant individual. She also described situations in which it appeared that squirrels emitted a bark out of frustration. Dollard et al. (1939) suggested that frustration is closely related to aggression. It appears, therefore, that the bark is an aggressive Response of red squirrel While predator present: Quiet Quiet After predator Predator left: None Bark Goshawk 4 Red-tailed Hawk Unidentified hawk 3 Great Horned Owl Hawk Owl | Marten D Total l 9 Rapid Bark Bark Bark squeak Bark and Quiet Chir 2 Bark 2 l 1 3 I 3 l 2) *Includes observations from Klugh (1918), Krasnowski (1969), Klassen (unpublished), Wetmore (personal communication), and the present study. 1977 call of lesser intensity than the chir call rather than an alarm call as previously suggested. Field work was conducted through the Depart- ment of Wildlife and Fisheries, University of Alaska. I thank Steve MacLean, University of Alaska; John Kelsall, Canadian Wildlife Service; and Fred Zwickel and Martin McNicholl, University of Alberta, for critically reading this manuscript and offering their constructive advice and criticism. I also thank Tom Wetmore, University of Alaska, for relating his observations to me and allowing me to cite them. Literature Cited Balph, D. M. and D. F. Balph. 1966. Sound communica- tion of Uinta ground squirrels. Journal of Mammalogy 47: 440-450. Carl, E.A. 1971. Population control in arctic ground squirrels. Ecology 52: 395-413. De Vos, A. 1958. Summer observations on moose be- havior in Ontario. Journal of Mammalogy 39: 128. Dollard, J., N. E. Miller, L. W. Doob, O. H. Mowrer, and R.R. Sears. 1939. Frustration and aggression. Yale University Press, London. 209 pp. Embry, P. C. 1970. Vocal communication of the red squir- tel, Tamiasciurus hudsonicus. M.Sc. thesis, University of Montana, Missoula. 51 pp. King, A. 1955. Social behavior, social organization, and population dynamics in a black-tailed prairiedog town in the Black Hills of South Dakota. Contributions of the NOTES 189 Laboratory of Vertebrate Biology, University of Michi- gan 67. 123 pp. Klugh, A.B. 1918. The behaviour of the red squirrel. Ottawa Naturalist 1: 9-12. Klugh, A. B. 1927. Ecology of the red squirrel. Journal of Mammalogy 1: 1-32. Krasnowski, P. V. 1969. Aspects of red squirrel (Tam- iasciurus hudsonicus) population ecology in interior Alaska. M.Sc. thesis, University of Alaska, College. 63 pp. Markham, O. D. and F. W. Whicker. 1973. Notes on the behavior of the pika (Ochotona princeps) in captivity. American Midland Naturalist 89: 192-199. Melchior, H. R. 1971. Characteristics of the arctic ground squirrel alarm calls. Oecologia 7: 184-190. Nodler, F. A. 1973. Food habits, vocalizations, and ter- ritoriality of Alaskan red squirrels (G. Tamiasciurus). M.Sc. thesis, University of Alaska, College. 86 pp. Searing, G. F. 1975. Aggressive behavior and population regulation of red squirrels (Tamiasciurus hudsonicus) in interior Alaska. M.Sc. thesis, University of Alaska, College. 99 pp. Smith, C. C. 1963. Territorial behavior in the genus of squirrel Tamigsciurus. M.Sc. thesis, University of Wash- ington, Seattle. 68 pp. Smith, C. C. 1968. The adaptive nature of social organi- zation in the genus of three [sic] squirrels Tamiasciurus. Ecological Monographs 38: 31-63. Received 9 November 1976 Accepted 19 January 1977 Predation on Nesting Glaucous-winged Gulls by River Otter RoBert G. Foottit! and Rosert W. BUTLER? ‘Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia V5A 1S6 22902 Brookridge Drive, North Vancouver, British Columbia V7R 3A8 Although there is evidence that birds constitute a small portion of the diet of the river otter, few cases of predation upon nesting seabirds have been recorded (Harris 1968). During the summers of 1967, 1970, and 1973 direct and indirect evidence of predation on nesting Glaucous-winged Gulls (Larus glaucescens) was observed on Mitlenatch Island, British Columbia (49°57’ N, 125°00’ W). A description of the island is given by Butler (1974). The river otter (Lutra canadensis) is an abundant animal along the coastal shorelines and offshore archipelagos of British Columbia (Cowan and Guiguet 1965). Since 1963, naturalists have lived on the island during the summer months and have observed river otters each year. Usually, individual or pairs of adult otters were sighted. In 1966, 1968, 1970, and 1973, however, young otters were also observed. During most summers only between 2 and 10 sightings of otters were made and only rarely were otters seen in the gull colony. Occasionally, single gull carcasses were found that appeared to have been dismembered. An actual observation of river otter predation was made in 1967 (Kennedy 1968). On 7 August an adult otter entered the gull colony, caught a young Glaucous-winged Gull by the back of the neck, and dragged the bird, its wings still flapping, into some dense bushes. In 1970 and 1973 additional evidence of apparent predation by river otter was found. On 30 June 1970 eight partially eaten adult gulls were found in a heavily trampled area among dense shrub, about 4.5 m (15 ft) in diameter. Otter feces were present and a trampled path led to the water. 190 During July otters were seen almost daily in the gull colony. Most sightings occurred between 1600 hours and sunset. Excited gulls were often heard at night, perhaps indicating continued otter activity. On | August two otters were observed eating a freshly killed gull hatchling. Twelve recently killed young gulls were found on 3 August in another area of approximately 36 m2 (400 ft?). During 1973 an otter was observed in the seabird colony as early as 6 June and gull carcasses were found by 9 June. On 30 July we saw an otter in the colony killing a week-old chick. Most dead gulls had their legs removed and teeth marks were evident in the head region. Most of the gulls were not eaten, indicating that surplus killing was taking place (Kruuk 1972). In 1973 some areas of the colony that afforded easy access from the water incurred extensive predation. On “F” Island, a small sub-island of 1.9 ha connected to the main island at low tide, approximately 250 Glaucous-winged Gulls were banded each summer, until 1973. On 27 July 1973 only two birds were banded, although a previous nest count of 26 June revealed 286 active nests. During this time interval dead young gulls were found scattered over “F” Island, all showing signs of otter predation. A recheck on 7 August again located only the two previously banded birds. On the “East Hill” area of the main island, only 60 young gulls were banded in 1973, compared to approximately 350 banded annually in previous years. Again, prebanding nest counts indicated little change in the number of active nests from previous years. Other areas of the colony experienced predation to a lesser degree. The steep cliffs along the south shore of the island were visited by predators where access from the water was available through crevices in the rocks. Although some of this evidence is circumstantial, it does indicate that the river otter is the probable predator of nesting Glaucous-winged Gulls. No other Prey Utilized by Urban Merlins L. W. OLIPHANT and S. MCTAGGART THE CANADIAN FIELD-NATURALIST Vol. 91 predators, such as mink (Mustela vison), have been observed on the island. During the summer of 1974, similar observations were made on Colville Island in Washington State (Hayward et al. 1975). On several occasions a single river otter was observed taking Glaucous-winged Gull chicks. The remains of adult birds, some dismembered, with their viscera eaten away, were found. The fact that uneaten gull carcasses were found on Mitlenatch Island makes it difficult to say to what extent the Glaucous-winged Gull makes up the summer diet of the river otter. As the gull colony on the island consists of approximately 2800 breeding pairs (Butler 1974), predation by the river otter would probably not seriously affect the colony size, unless a great deal of surplus killing occurred over an extended period of time, or an increased number of otters utilized this potential summer food source. We are grateful for the helpful criticisms of R. Wayne Campbell and N. Verbeek. Literature Cited Butler, R. W. 1974. The feeding ecology of the North- western Crow on Mitlenatch Island, British Columbia. Canadian Field-Naturalist 88: 313-316. Cowan, I. McT. and C. J. Guiguet. 1965. The mammals of British Columbia. British Columbia Provincial Muse- um Handbook 11: 1-414. Harris, C. J. 1968. Otters: A study of the recent Lutrinae. Weidenfeld and Nicolson, London. 397 pp. Hayward, J.L. Jr., C.J. Amlaner, W.H. Gillet, and J. F. Stout. 1975. Predation on nesting gulls by a river otter in Washington State. Murrelet 56: 9-10. Kennedy, K. 1968. River otter feeding on Glaucous-winged Gull. Blue Jay 26: 109. Kruuk, H. 1972. The urge to kill. New Scientist 54: 735-737. Received 6 February 1976 Accepted 23 December 1976 Department of Veterinary Anatomy, Western College of Veterinary Medicine, University of Saskatchewan, Saskatoon, Saskatchewan S7N 0WO Merlins (Falco columbarius) have nested regularly in the city of Saskatoon, Saskatchewan, since 1971. In 1974, an attempt was made at one nest site to determine the species composition of prey taken. From this study, it was estimated that 90% of the prey consisted of House Sparrows (Passer domesticus), the most abundant small bird in the area (Oliphant 1974). A more detailed study of prey utilization was 1977 undertaken at two urban nests during 1975 to supplement the data gathered the previous year and to allow comparisons with observations made on prairie-nesting Merlins. Methods During the breeding season, prey remains, con- sisting primarily of flight feathers, feet and heads, were collected from under plucking perches near nest sites. The University of Saskatchewan Biology Department study skin collection was used to confirm identification. Sometimes prey was identified by direct observation with binoculars as it was brought to the nest site by the male or as it was being eaten by the female or fledglings. Observations and collection of remains were made onan almost daily basis from mid- June through mid-July in 1974. During 1975, two nest sites were visited nearly every day during May and June. Occasional observations and collection of prey remains were also made during March and April. Both of the nest sites (one site studied in both years) were situated adjacent to residential areas with open park-like areas nearby. The nest site studied both years was located on the University of Saskatchewan campus. The hunting territory of the male included adjacent residential areas and the agricultural plots and livestock areas of the university. The other nest was in a cemetery in town. The hunting territory of this male included the cemetery and adjacent residential and semi-industrial areas. Limited data was also collected on prey taken by urban Merlins outside the breeding season. This included observations on immature Merlins soon after they had become independent of their parents (including two groups of young fledged from an artificial nest), wintering Merlins, and observations on the hunting success of a single trained Merlin. Results and Discussion During the breeding season, nearly all hunting is done by the male, and birds were the only recorded prey species. Of the total sample of prey, 8% could not be identified for certain. These were all sparrow-sized birds observed through binoculars only. The majority are presumed to have been House Sparrows. House Sparrows accounted for 69% of the 162 prey items that were definitely identified. Three other species, Horned Lark (Eremophila alpestris), Bo- hemian Waxwing (Bombycilla garrula), and Robin (Turdus migratorius) accounted for an additional 14%. The remaining 15 species (17%) were recorded less than 5 times each (Table 1). The prey species were not utilized randomly, primarily because most of the species are not uniformly available throughout the breeding season. NOTES 19] TABLE |—Prey species utilized by urban nesting Merlins Number of 4% of Species individuals total House Sparrow (Passer domesticus) 112 63.6 Unidentified sparrow-sized birds thought to be Passer domesticus 14 8.0 Horned Lark (Eremophila alpestris) 10 Soll Bohemian Waxwing (Bombycilla garrula) 7 4.0 Robin (Turdus migratorius) 6 3.4 Cedar Waxwing (Bombycilla cedrorum) 4 D8} Dark-eyed Junco (Junco hyemalis) 3 1.7 Swainson’s Thrush (Aylocichla ustulata) 2} 1.1 Ruby-crowned Kinglet (Regulus calendula) 2 1.1 Red-eyed Vireo (Vireo olivaceus) ” 1.1 Song Sparrow (Melospiza melodia) 2 lel Lapland Longspur (Calcarius lapponicus) yD, 1.1 Chestnut-collared Longspur (Calcarius ornatus) 2 1.1 Tree Swallow (Urodoprocne bicolor) 2 1.1 Flycatcher (Empidonax sp.) ] Hermit Thrush (HAylocichla guttata) 1 0.6 Western Meadowlark (Sturnella neglecta) l 0.6 Savannah Sparrow (Passerculus sandwichensis) l 0.6 White-throated Sparrow (Zonotrichia albicollis) ] 0.6 Lincoln’s Sparrow (Melospiza lincolnii) ] 0.6 Total 176 Bohemian Waxwings were recorded only in March, April, and early May. This species generally leaves the Saskatoon area by early May. Only two species, Bohemian Waxwings and House Sparrows, were recorded during April. The diversity of prey species was greatest in May with 15 species recorded. This coincides with the height of passerine migration in the Saskatoon area. In contrast, only eight species were recorded in June with three species (House Sparrow, Horned Lark, and Robin) making up over 90% of the prey items. The percentage of House Sparrows taken steadily increased from 40% in April to about 80% in 192 June and July. This tendency may account for the very high percentage of sparrows recorded in 1974 since prey items were only recorded from mid-June through July. Although not recorded as a definite prey item, Common Flickers (Colaptes auratus) were unsuc- cessfully pursued on at least two occasions and old remains of this species were found near one nest. Lawrence (1949) found remains of a Flicker in the nest of a Merlin in Ontario and non-breeding Merlins have been recorded pursuing Flickers (Fox 1964; W. Harris, personal communication). The prey species utilized by urban nesting Merlins differ considerably from those reported for prairie- nesting individuals of the same subspecies (richard- sonii). Remains found in nests near Kindersley, Saskatchewan, consisted of 53.5% Horned Larks, 13.6% Chestnut-collared Longspur, 13.3% Cowbird, and 20.1% native sparrows (Fox 1964). Similarly, Hodson (1976) found that Horned Larks accounted for 50% of the prey, Chestnut-collared Longspurs 37%, native sparrows 2%, blackbirds 4%, and other species (including unidentified) 7%. This was com- puted from 2070 prey remains collected at nest sites in southern Alberta. Certainly the most obvious factor in accounting for the differences in prey utilization between urban and prairie-nesting Méerlins is the different relative abundances of available prey in the two environ- ments. The prey recorded in this study indicate that Merlins take advantage of species that happen to be abundant in the area at a given time. Certain species, however, such as Clay-colored Sparrows, Spizella pallida, and various warblers were not recorded as prey although they are more common than most of the prey species recorded with the exception of House Sparrows. Whether this was due mainly to the low sample size or whether certain birds are less likely to be preyed upon than others is not clear. Hodson (1976) has noted that certain abundant potential prey species are rarely taken. He has suggested that the specific habitat preference (grazed versus undisturbed grassland) and the activity pattern of the feeding birds account for the heavy utilization of certain species rather than others of equal or greater abundance. Birds preferring undisturbed grasslands and having less active feeding patterns were relatively safe from Merlin predation. Similar factors may be involved in affecting prey vulnerability in the urban environment. Merlins wintering in Saskatoon prey heavily on the few species available. Bohemian Waxwings and House Sparrows are the most common prey species utilized during the winter. This is also true of the THE CANADIAN FIELD-NATURALIST Vol. 91 Merlins wintering in Edmonton, Alberta (R. Fyfe in Trimble 1975). The only other species definitely recorded during the winter months was a single Dark- eyed Junco, Junco hyemalis. There is much evidence to suggest that large insects are utilized when abundant, especially by immature birds (see Trimble 1975). Insects may often be the first prey taken by newly fledged Merlins. Hodson (1976) cites an observation of a family of newly fledged young “pursuing and eating grasshoppers during a heavy hatch of these insects.” Immature Merlins have been observed capturing dragonflies on the wing in the Saskatoon area in late summer (Oliphant 1974). Young Merlins fledged from an artificial nest structure in Saskatoon during 1975 were observed feeding extensively on dragonflies, which are often abundant in late summer. These birds were also seen pursuing House Sparrows but all the attempts observed were unsuccessful (Oliphant and Thomp- son, unpublished data). Observations of a trained, immature female Merlin indicated that dragonflies were much more easily captured than birds although they provided relatively little biomass per kill. Approximately 50% of all attempts at capturing dragonflies were successful during the first two months after fledging; success at capturing birds was less than 5% during the same period. Dragonflies and other large flying insects present in abundance during the first 1-2 months after fledging may be extremely important in sustaining young Merlins as they develop their flying skills. This study was supported in part by a grant (#A9886) from the National Research Council of Canada. The authors thank W. J. P. Thompson for help in gathering prey remains. Literature Cited Fox, G. A. 1964. Notes on the western race of the Pigeon Hawk. Blue Jay 22: 140-147. Hodson, K. 1976. The ecology of Richardson’s Merlins on the Canadian Prairies. M.Sc. thesis, University of British Columbia. Lawrence, L. de K. 1949. Notes on nesting Pigeon Hawks at Pimsi Bay, Ontario. Wilson Bulletin 61: 15-25. Oliphant, L. W. 1974. Merlins — the Saskatoon falcons. Blue Jay 32(3): 140-147. Trimble, S. A. 1975. Habitat management series for unique or endangered species. Report Number 15, Merlin, Falco columbarius. United States Department of the Interior, Bureau of Land Management. Received 13 August 1976 Accepted 30 January 1977 oa NOTES 193 Use of Man-made Structures as Nest Sites by Pigeon Guillemots R. WAYNE CAMPBELL British Columbia Provincial Museum, Victoria, British Columbia V8W I1A1 In western coastal North America, the Pigeon Guillemot (Cepphus columba) nests typically in natural crevices in a variety of habitats. Drent et al. (1964) and Campbell (1975) have described six major site-types of enclosed nests on the British Columbia coast. In addition, Dawson and Bowles (1909), Thoresen and Booth (1958), Bowman (1961), and Drent et al. (1964) reported that open ledge sites are occasionally used. Observations during the summers of 1970-1976 revealed the use of man-made struc- tures for nest sites by Pigeon Guillemots. These are listed in Table 1. Only two of the nine nest sites listed were actually checked. Breeding evidence for the remainder con- sisted of seeing adults carrying fish, presumably for chicks, or adults flying to nest sites. Nests are suspected, but not confirmed, for sites at Namu and Campbell River. Eight sites were associated with wharves. The eggs at Sandspit (Queen Charlotte Islands) and Chemainus (Vancouver Island) were deposited on narrow ledges under the wharves. In all cases vertical planks prevented the egg(s) from rolling into the water. Occasionally, adults were observed swimming and/or “bill-dipping” under the wharves and on one occasion I watched an adult land, with some difficulty, on its nest ledge. At Horseshoe Bay, near the government ferry terminal, an adult Pigeon Guillemot, carrying a fish, was seen entering a group of wooden pilings. As seen from the ferry, the nest site appeared to be on top ofa broken timber in the center of the pilings. Although most of these man-made structures used by guillemots for nesting have been constructed within the past 30 years or so, it is not known when such use first began. Available habitat probably does not limit numbers of Pigeon Guillemot in British Columbia as it does for Black Guillemot (Cephhus grylle) in northern Alaska (Divoky et al. 1974). The use of man-made structures, however, supports Storer’s (1952) view that guille- mots are plastic in their choice of nest sites and that cover appears to be a “principal requirement” for nesting. I thank S. G. Sealy for comments on an earlier draft of this manuscript. Literature Cited Bowman, R. I. 1961. Late spring observations on birds of South Farallon Island, California. Condor 63: 410-416. TABLE 1—Nesting data for Pigeon Guillemots using man-made structures in British Columbia, listed by localities from north to south Type of Locality Date structure Prince Rupert 5 July 1976 Wharf Sandspit 16 June 1974 Wharf Namu 20 June 1976 Wharf Port Hardy 1 July 1975 Wharf Campbell River 10 June 1976 Wharf Ucluelet 24 July 1970 Wharf Horseshoe Bay 6 July 1974 Piling Chemainus 22 June 1974 Wharf Sydney 25 June 1974 Wharf Number of Breeding breeding pairs evidence 3+ Adults flying to nest ledge I+ One egg; two eggs 1 Adult swimming under wharf! I+ Adult flying to nest ledge 3+ Adults on water nearby! 1 Adult carrying fish 1 Adult carrying fish 2+ Two eggs; two eggs 1 Adult flying to nest ledge 'Not positive, but supportive. 194 Campbell, R. W. 1975. Seabird colonies in Skidegate Inlet, Queen Charlotte Islands, British Columbia. Syesis 8: 355-361. Dawson, W.L. and J.H. Bowles. 1909. The birds of Washington State. Occidental Publishing Company, Seattle, Washington. 767 pp. Divoky, G. J.. G. E. Watson, and J. C. Bartonek. 1974. Breeding of the Black Guillemot in Northern Alaska. Condor 76: 339-343. Drent, R. H., G. F. van Tets, F. Tompa, and K. Vermeer. 1964. The breeding birds of Mandarte Island, British THE CANADIAN FIELD-NATURALIST Vol. 91 Columbia. Canadian Field-Naturalist 78: 208-263. Storer, R. W. 1952. A comparison of variation, behavior, and evolution in the seabird genera Uria and Cepphus. University of California Publications in Zoology 52: 121-222. Thoresen, A. C. and E. S. Booth. 1958. Breeding activities of the Pigeon Guillemot (Cepphus columba). Walla Wallac College Publication Number 23: 1-37. Received 16 September 1976 Accepted 9 November 1976 Lysurus gardneri, an Uncommon Stinkhorn Observed in Eastern Ontario VINCENT NEALIS,! J. GINNS,2 and W. I. ILLMAN! ‘ELBA, Department of Biology, Carleton University, Ottawa, Ontario KIS 5B6 2Biosystematics Research Institute, Canada Agriculture, Ottawa, Ontario KIA 0C6 Stinkhorn is the vernacular name applied to a small group of fungi (Basidiomycetes: Gasteromycetes: Phallales). The name is derived from the general shape, like a cow’s horn, and the fetid odor of the mature state. The discovery of an unusual species, Lysurus gardneri Berk. (family Clathraceae), was made in 1975 at Navan, Ontario (near Ottawa). The only prior report of this species in Canada is from British Columbia by M. C. Melburn (1966. Victoria Naturalist 22(5): 49). Additional records in the National Mycological Herbarium (DAOM) at the Central Experimental Farm in Ottawa are one collection, each, from Ottawa, nearby Russell, Ontario, and Papineau County, Quebec (also near Ottawa). We have learned of a collection on cow dung near Tweed, Hastings County, Ontario, made on 4 October 1976 by F. van Gerwin and deposited in the herbarium of the University of Toronto as TRTC 47654. These Canadian records extend the published North American range of the species, which has been reported from several northeastern states of USA, to as far west as Ohio, and also from California. It is known from several countries of western Europe, South America, and Australasia (from New Zealand to India). Conjectures in the literature are that the species is introduced from the tropics or south temperate regions; it represents the sole representative of the family Clathraceae to be reported in Canada, our other stinkhorns being members of the Phal- laceae (with the spore-containing gleba covering an apical portion of the stem or an undivided cap which hangs down around the stem from its tip). The habitat of L. gardneri, from data with the DAOM collections, was relatively undecomposed horse manure in gardens, greenhouses, and piles of stable-cleanings. The Navan specimens (DAOM 154172) were restricted to a narrow strip of land ina vegetable garden which had been a tethering line for ponies. The following observations were made from DAOM 154172. The young stage (Figures 1-3) of the stinkhorn is a white soft but leathery globe, commonly called an “egg”; this arises from white mycelial strands. The egg, recognizable when only a few millimetres in diameter, may enlarge to nearly 5 cm. In the early stages (Figure 2), the white fleshy shell (or volva) encloses a layer of gray gelatinous matter (also part of the volva). Inside these layers is a brown tissue from which finger-like projections develop, ap- parently at the expense of the brown tissue. By the time the volva ruptures, both the gray gelatin and the brown matter are almost gone. Exposed to the air, the remaining gray matter rapidly disappears. Develop- ment now proceeds with the elongation of the stem, the growth and separation of the fingers, and the differentiation of a brown sporulating gleba on the wrinkled surface of the fingers (Figures 3 and 4). At full maturity the white hollow stem is 10 to 15 cm long and about 3 cm in diameter, and the pale orange- yellow fingers (five to seven in number) are about 3 cm long and support a brown spore-containing fetid exudate that is conspicuous and attractive to flies. 1977 NOTES 195 Ficures 1-4. Lysurus gardneri(DAOM 154172). |. Four immature stinkhorns in the egg stage. Two have ruptured exposing the apex of the pileus. 2. Vertical sections through five eggs. The youngest egg (top) shows little differentiation, being composed of the brown primordial tissue and the gelatinous outer layer. The other eggs, with the most mature at the bottom, show an increase in egg size as development of the fingers and stalk progresses. 3. Eggs and fully mature stinkhorns. 4. Mature stinkhorn with fetid spore mass almost completely gone from the rugose surface of the fingers. Scale: pen length in Figure | is about 14 cm. 196 Several other observations made on the Navan collection relate to the life cycle and means of dispersal of the stinkhorn. The manure substrate had been fresh in the spring of 1975. A mature stinkhorn was first noted on 12 August, about 3 months later. The fungus probably overwintered in the mycelial state in the garden soil and the fresh manure provided the necessary nutrition to induce fruiting. Alter- natively, but seemingly less likely, it is possible that overwintered stinkhorn spores may have been deposited by insect vectors on the relatively fresh manure, the entire development of the fungus, from spore to maturity, taking less than 3 months. The garden was cleared of vegetables by 17 September and, despite its exposed position, L. gardneri flourished. In fact, unlike many large fleshy fungi, the stinkhorn seemed to prosper during hot THE CANADIAN FIELD-NATURALIST Vol. 91 sunny weather. Clusters of eggs, which had persisted relatively unchanged throughout the cool moist period of 17 to 26 September, produced mature stinkhorns after only 2 days of warmer dry weather. The maturation of this fragile, quite temporary fruiting state is apparently adapted to the stinkhorn’s dependence on insects to disperse the spores; flies were noticeably inactive during the cool moist weather. Besides the numerous green bottle flies (Calli- phoridae), two types of beetle were noted, some small rove beetles (Staphylinidae) and a single dung beetle (Ataenius strigatus) (Scarabaeidae). Stinkhorns were still numerous when the final observations were made on 31 September. Received 26 October 1976 Accepted 7 January 1977 News and Comment Editor’s Report The numbers of manuscripts received and accepted for publication in The Canadian Field- Naturalist in recent years are shown in Table |. Note that the final disposition of a manuscript may or may not be determined in the same year in which the paper 1s submitted. TABLE | Number of manuscripts % Year received accepted accepted 1973 153 117 76 1974 152 115 76 1975 167 118 71 1976 147 _ a Table 2 categorizes, according to fields of study, the manuscripts published in The Canadien Field- Naturalist during the past three years. The papers published in issues Number |, 2, and 4 of 1976 plus issue Number 4 of 1975 (four issues published per year) have been further categorized in Table 3. If this summary is used as the basis for future comparisons of the proportions of papers submitted or published on the different taxonomic groupings of organisms, new trends or emphasis should become apparent. Request for Participants International Shorebird Surveys, 1977-78 A cooperative International Shorebird Survey scheme was started in 1975 to obtain information on shorebird migration and to identify and document areas of major importance. This scheme has been highly successful, with much very valuable informa- tion on shorebird distribution and migration coming from contributors throughout eastern Canada and the USA, the Caribbean Islands and Central and South America. Information from the scheme will be valuable in assessing requirements for the future protection and conservation of the birds and their habitat. In 1977 we are anxious to continue and extend the scheme in as many areas as possible. Any TABLE 2 Number of manuscripts published in one year Major subject 1974! 1975! 19762 Birds 56 39 42 Mammals 21 22 33 Plants 16 13 21 Others 14 16 28 Total 107 90 124 "Includes issues Number I, 2, and 3 plus issue Number 4 from the previous year’s volume. *Includes issues Number |, 2, and 4 plus issue Number 4 from the previous year’s volume. The special raptor issue, Number 3, which contained 8 articles and 6 notes on birds, is not included. TABLE 3 Number of manuscripts Major subject Articles Notes Birds 6 36 Mammals 12 21 Plants 6 15 Fishes 2 8 Amphibians & reptiles l 6 Invertebrates 4 3 Others 3 I Total 34 90 observer who may be able to participate in regular survey counts of shorebirds during spring and autumn migration periods, as well as during the winter in shorebird wintering areas, are asked to contact one of the undersigned. Occasional counts from observers visiting shorebird areas on an irregular basis would also be most welcome. For areas in Canada: Dr. R.I.G. Morrison, Canadian Wildlife Service, 2721 Highway 31, Ottawa, Ontario, Canada KIA OE7 For areas in the USA, Caribbean Islands, Central and South America: Brian A. Harrington, Manomet Bird Observatory, Manomet, Massachusetts, USA 02345. 197 198 Thank You Earl Godfrey The recent retirement of Dr. W. Earl Godfrey, Associate Editor (Ornithology) of The Canadian Field- Naturalist from 1947 to 1976, from the Editorial Board of The Canadian Field-Naturalist cannot pass without formal recognition. For his almost thirty years of dedicated service, on my own behalf and that of former Editors of the journal who have had the pleasure of working with Earl, I thank him most sincerely. This acknowledgment is not only for his willingness to serve as a referee for manuscripts in ornithology, the largest single field of journal papers, but also for his devotion to a task requiring someone with his extensive knowledge. Only some of us realize the amount of time and effort Earl has expended to review and edit manuscripts. In spite of his broad expertise in nomenclature, distribution, plumages, behavior, and ecology of birds, many hours were often taken up in the meticulous study of past records, files, manu- scripts, references, and the examination of bird specimens so that he could gain fresh insight into a Request for Information Shorebird Color-marking In 1977, the Canadian Wildlife Service will again be carrying out extensive banding and color-marking of shorebirds in James Bay. Last year, over 12 400 shorebirds were captured during July and August resulting in over 580 reports of color-marked birds in eastern North America and South America. Much valuable information on migration routes is being obtained and observers are again asked to look out for and report any color-dyed or color-banded shore- birds that they may see. Reports should include details of species (with age if possible), place, date, color- Request for Information Color-banded Semipalmated and Least Sandpipers Last year the Surinam Forest Service color-banded nearly 3300 Semipalmated and Least Sandpipers, resulting in 14 spring and summer sightings and recoveries from the United States and Canada. In 1977 again large numbers of these species will be color-banded along the Surinam coast. As in 1976, birds will be banded above the tarsus (“knee”) witha standard aluminum band and TWO ORANGE plastic bands of about the same size as the aluminum THE CANADIAN FIELD-NATURALIST Vol. 91 subject before he manuscript. Earl has always been an active field person, a staunch supporter of The Canadian Field- Naturalist, and a strong believer in the journal’s worth. He must be commended for the particular attention and encouragement he has given to amateurs. I have been especially appreciative of his concern for, and insistence upon, accuracy, and for his regard for the proper use of the English language. Moreover, although his comments show perception and concern, they are also kind and constructive. His retirement from the post of Associate Editor of The Canadian Field-Naturalist fortunately does not mean that he will no longer be willing to review manuscripts on subjects of special interest to him or that would profit from his particular judgment and experience. Indeed, I intend to call on him from time to time in the future. interpreted and evaluated a LORRAINE C. SMITH, Editor marks and, if possible, notes on the numbers of other shorebirds present. For color-dyed birds, please record the color and area of the bird that was dyed. For color bands and standard metal leg bands, please record which leg the bands were on, whether they were above or below the “knee,” the colors involved, and the relative position of the bands if more than one was on a leg (e.g., right leg, blue over metal, etc.). All reports will be acknowledged and should be sent to Dr. R. I. G. Morrison, Canadian Wildlife Service, 2721 Highway 31, Ottawa, Ontario, Canada KIA OE7. band. We again ask birders to look out for these birds and to send reports of observations to Arie L. Spaans, Surinam Forest Service, P.O. Box 436, Paramaribo, Surinam, South America. Please report species, date and location of observation, the position of the aluminum and color-bands—left or right leg, and, if more than one band is on a leg, which band is above, which below, and which in the middle (some birds have all three bands on one leg)—and the number of color-banded birds involved. 1977 NEWS AND COMMENT 199 Pilot Study for a Biological Survey of the Insects of Canada Have you received a questionnaire? The Entomological Society of Canada has been awarded a contract by the Canadian Government to conduct a Pilot Study for a Biological Survey of the Insects of Canada. This project is intended to establish the foundations for a continuing biological survey, and a major aim is to assess the resources and needs of Canadian research in the identification, distribution and biology of insects (including arachnids and other related forms). Four types of questionnaire have therefore been distributed. If any readers have information to contribute on relevant resources or needs and were overlooked in the original mailing, we would be very pleased to receive requests for the applicable question- naires as listed below. (1) Questionnaire to individual entomologists, to Colonial Waterbird Group First Annual Meeting The Colonial Waterbird Group, organized during the Wading Bird Conference at Charleston, South Carolina last October 1976, will hold its first annual meeting on 21-23 October 1977, at Northern Illinois University, in DeKalb. The conference will include paper sessions, subgroup meetings (surveys, conser- vation, etc,) and an important business session. Any person wishing to present a paper on an aspect of research or management of pelicans, cormorants, ascertain the location of personnel and programs, and seek information on the state of knowledge in Canada of their taxonomic or ecological groups of interest. (2) To resource managers, environmentalists and other users of information on insects, to ascertain their present and future needs for entomological information. (3) To directors of institutions conducting entomo- logical research in Canada, to ascertain programs and facilities. (4) To curators of collections in Canada and else- where, to ascertain the whereabouts of significant holdings of Canadian arthropod material. Secretariat, Biological Survey Project, 202-1316 Carling Avenue, Ottawa, Ontario KIZ 7L1 herons, ibises, gulls, terns, alcids or other colonial waterbirds should submit a single page abstract no later than 15 August 1977 to the National Audubon Research Department, 115 Indian Mound Trail, Tavernier, Florida 33070. Additional information on the conference will appear in the mid-summer CWG newsletter, or may be obtained by writing the above address. Book Reviews ZOOLOGY Biology of the Kaminuriak Population of Barren-ground Caribou. Part 2 By Frank L. Miller. 1974. Canadian Wildlife Service. Ottawa, Report Series Number 31. 88 pp. $3.00. The Canadian Wildlife Service in 1966 began a 2'4- year intensive study of the Kaminuriak population of barren-ground caribou. The study was divided into four parts, each the responsibility of the CWS biologist: total numbers, distribution, recruitment and mortality; sex and age composition; seasonal physical and reproductive condition; and winter range evaluation. Although the study was undertaken primarily by the Canadian Wildlife Service, the game agencies of Manitoba, Saskatchewan, Alberta, and the Northwest Territories all contributed personnel at various stages of the project. The results of the research project were reported to the Administrative and Technical Committees for Caribou Preservation in 1970. This publication constitutes a report of the second part of the study. (Part | by G. R. Parker, which included total numbers, mortality, recruitment, and seasonal distribution was published as CWS Report Series Number 20, in 1972. Reviewed in The Canadian Field- Naturalist 88(3): 376-377.) The current report is divided into two parts: (1) dentition as an indicator of age and sex, (2) socialization and population analysis. A sample of 999 caribou, including all important age and sexual classes, was collected systematically at four seasonal periods from March 1966 to July 1968. Nine hundred and forty-three caribou were collected from the Kaminuriak population and 58 from the adjacent Beverly population. This sample represented approxi- mately 1.5% of the total population (63,000, Parker 1972). This number may have provided a statistically valid population sample, but it is large enough to raise concerns among conservationalists as well. It is to be hoped that other caribou populations need not be subjected to such intensive sampling. Age was estimated by tooth eruption and replace- ment, by linear tooth measurement, and by micro- scopic histological examination of the cementum layer of mandibular teeth. The author presented a convincing case for the annual deposition of an annulus (winter rest line). He reported the eruption of the permanent dentition between the ages of 24 and 29 months. The difference between Miller’s conclusion and my earlier work (Preliminary investigation of the barren-ground caribou. Part 2. C.W.S. Wildlife Management Bulletin Number 10b, 1954) is the result of my reliance upon a small control sample of 12 mandibles from tagged reindeer of known age from the Mackenzie Delta herd. Considerable variation in tooth eruption at standard ages was noted as well as the need for more data on differential wear of forest and tundra caribou relative to differing feeding behavior. If one also added genetic differences between reindeer and caribou populations, it might be possible to account for the differences in conclusions. The subsequent discussion of cohort mortality could have been clarified if the author had distinguished density-dependant and density-independant mor- tality factors. The second part on socialization commences witha definition of the various types of age and sexual groupings which the author defines as bands. The statistical analysis of the occurrence of radio-tagged caribou indicated the social cohesion of these bands. Miller postulated that the winter groupings are basic, and the post-calving aggregation serves to reunite the previously segregated winter bands. He further postulated that the herding behavior of caribou evolved as a defense against wolf predation, because the herds saturate the wolves’ hunting territories. He might have considered the migratory behavior of wolves in packs over much of the year. which would counteract that defense mechanism to a considerable degree. In my view, the long caribou migration from the winter ranges in spring is an effective way of leaving behind the wolves which commence to den in April-May. Fawning then takes place in areas where the resident wolves are restricted to their denning territories. The final section on population analysis of the Kaminuriak population was based upon the age and sex composition of the sample population. It clearly indicated a weak 1962 cohort such as is frequently found in fish population studies. Survival curves were prepared by adjusting the numbers for this weak class, and significantly different male and female survival rates were indicated. In his conclusion the author recognized the difficulties in overcoming sampling bias on various age and sex classes. A. W. F. BANFIELD Institute of Urban and Environmental Studies, Brock University, St. Catharines, Ontario L2S 3Al 200 1977 BOOK REVIEWS 201 Biology of the Kaminuriak Population of Barren-ground Caribou. Part 3 By Donald R. Miller. 1976. Canadian Wildlife Service, Ottawa, Report Series Number 36. 42 pp. $1.75. This publication which is subtitled “Taiga winter range relationships and diet,” is a report on the last part of the intensive study of the Kaminuriak popu- lation. T. C. Dauphine, Jr. willsoon publish as Part 4, the report on growth, reproduction, and nutritional condition (see New Titles list in The Canadian Field- Naturalist 91(1)). The author studied the winter range and diet of the Kaminuriak population in northwestern Manitoba and northeastern Saskatchewan by means of vegeta- tion quadrats, exclosures, and the study of feeding activity, feeding craters, and rumen contents. The potential forage was determined by quantitative data on standing crop and percentage plant cover. Terres- trial lichens covered 50 to 90 percent of the ground in exclosure plots and the standing crop of terrestrial lichens varied from 2000 to 7000 kg dry weight per hectare. Lichen regeneration of primary thalli on artifically denuded plots occurred on all plots after three growing seasons. The author found no relation- ship in the terrestrial lichen standing crop with age of the stand over 30 years, although the data in Figure 4 appear to suggest increasing biomass in similar stands up to 120 years of age. Climatic factors, especially snow depth and crust hardness, were reported to account for the sudden changes observed in caribou diet. The diet of early winter was predominantly terrestrial lichens. This changed to arboreal lichens and woody browse in late winter. As the snow melted in spring the caribou fed heavily on exposed lichens and higher plants along migration routes. Forest fires were considered bene- ficial because they provided heterogeneity of plant cover in the taiga. Based upon quantitative caribou food studies elsewhere, the winter range of the Kaminuriak population was estimated to have the carrying capacity for 360 000 caribou (about 6 times the current population). Unfortunately, there were several errors in the report, including Figure 4, and a sheet of “Errata” has been issued. Readers should ensure that the page is included in their copy of the report. With the publication of these reports, the Kaminuriak popula- tion has become the most intensively studied of all the caribou “herds,” and our understanding of its popula- tion dynamics has been placed ona level equivalent to that of other well studied species such as the African elephant. A. W. F. BANFIELD Institute of Urban and Environmental Studies, Brock University, St. Catharines, Ontario L2S 3A1 The Insects and Arachnids of Canada. Part 2: The Bark Beetles of Canada and Alaska By Donald E. Bright, Jr. 1976. Agriculture Canada, Ot- tawa. 241 pp., illus. $7.00 in Canada, $8.40 other countries. This book is part of a Canadian Faunal Series being developed by staff of the Biosystematics Research Institute, Canada Agriculture, as identification guides to the arthropods of Canada. These guides are designed to permit the identification of organisms by the general biologist, senior technician, or advanced amateur. Although Bark Beetles of Canada and Alaska is Part 2 of the series, it is the first to be published. Part 1, a handbook of entomological techniques, has been delayed by the preparation of illustrations. Other contributions in the series have been submitted for publication or are in late stages of preparation. This is the first comprehensive treatment of species of bark beetles in Canada since J. M. Swaine’s Canadian Bark Beetles, published in 1918. Swaine’s book provided excellent coverage of the bark beetles but has long been out of print and unavailable to most people. Consequently this handbook will once again provide students, amateur collectors, foresters, and others with a means of identifying various species of Scolytidae. The reader is given a brief account of the general biology of bark beetles which is followed by a discussion of the types of galleries created by the adult and larval forms. These types are illustrated by a number of photographs of typical bark beetle pat- terns. I would have preferred seeing these illustrations adjacent to the pertinent section in the text rather than at the back. Adult anatomy is described and illustrated in only sufficient detail to include the parts of the body used in the descriptions and in the keys. Descriptive ter- minology is further defined ina glossary at the back of the book. The user is not burdened with more descriptive morphology than is required to use the book for its intended purpose, 1.e., identification. 202 Two hundred and fourteen species in 45 genera known or suspected to occur in Canada and Alaska are dealt with. For each species the following information is given: synonomy (if any); a brief diagnosis emphasizing the most obvious or easily visible morphological features; host plants; geo- graphical distribution supported by a map of locali- ties where the species has been found; and a brief summary of the biology if known. A particularly valuable feature is the reference to more detailed accounts of biology of the species where known so anyone can proceed beyond mere identification if he so desires. The book is well organized and easy to use. Those not familiar with scolytid taxonomy should have no difficulty in keying to genera and then to species. Important diagnostic features of parts and entire species are illustrated by 173 photographs taken with A Book of Canadian Animals By Charles Paul May. Illustrations by John Crosby. 1976. Macmillan of Canada, Toronto. 115 pp. Paper $4.95. This book was written for, and should appeal to, children aged from about eight to twelve years. It contains twenty-eight short chapters, each describing a different mammal and some of its habits. For each chapter, a line drawing of the adult and young for the species concerned accompanies the text. The chapters all follow roughly the same format. The animal is introduced and the reader is told where in Canada it is usually found. Next we are told what it eats, where it lives (i.e., in a burrow or whatever), and any unusual habits are mentioned. A major focus for each chapter seems to be when and how many “babies” are born to mother and father animal, and for some, whether they make good pets. Most chapters finish with a brief passage relating the animal’s usefulness or special interest to man. I gave the book to a grade five teacher who circulated it amongst her pupils. Their responses to it were varied but all seemed to enjoy and like the book. Two consistent comments were that the chapters were much too short and gave the children a sense of incompleteness; and the children liked the drawings but wished they were in color and much larger. Generally, the children found the book enjoyable and readable, but would not exactly call it entertainment. Perhaps this was because they read it in school, but on a volunteer basis. In my opinion, a book of animals would contain not just mammals, as this one, but also birds and reptiles, or perhaps even insects and fish. But I do not believe THE CANADIAN FIELD-NATURALIST Vol. 91 the aid of a scanning electron microscope. For instance, dorsal and lateral views of a representative species of 38 of the 45 genera are included, as well as superb illustrations of antennae and tibiae. These figures are referred to in the pertinent sections of the keys with only one omission that I noted. Reference to Figures 133 and 171, Xyleborus, is omitted from couplet 34 of the generic key. This handbook provides a much needed reference guide to the Canadian bark beetles and I would recommend it for inclusion in university libraries, and as a personal choice for anyone interested in knowing a little more about this group of beetles. J. B. THOMAS Canadian Forestry Service, Sault Ste. Marie, Ontario P6A 5M7 the title is misleading to youngsters. The coverage of species is wide and interesting, with the omission of caribou, black bear, mountain sheep, and elk. Other species found here, including the pocket gopher, the star-nosed mole, and the pika, are not usually represented in animal surveys for children. The preoccupation in the book with “babies” of the animals discussed is presumably a technique for capturing and maintaining the interest of youngsters. The unfortunate cumulative impression is that the animals are “cutesy-pie” and “cuddly”; that they make good or bad pets, or that they are scary and dangerous when hungry (see the pages on the gray wolf, p. 109). This is the information that persists in the mind long after the more important details of life history have vanished. Is this the intent of the book? The problem of human values and attitudes pervades much of the wildlife literature and film meant for children and calls into question precisely what is the role of such media in pre-adult nature education. I suggest that the potential impact of this book goes beyond providing information in a simple manner. It extends into the realm of reinforcing a paternalistic and anthropocentric attitude toward nature. The technique is subliminal and the impact is cumulative. I would suggest that the use of this book by youngsters should be accompanied by careful guidance from parents and teachers. BRIAN WILKES 688A Winchester Avenue, Nanaimo, British Columbia VOR 4B8 1977 BOOK REVIEWS 203 An Investigation of Caribou Range on Southampton Island, N.W.T. By G. W. Parker. 1975. Canadian Wildlife Service, Ot- tawa, Report Series Number 33. 83 pp. $2.75. Excessive hunting on Southampton Island in northern Hudson Bay led to the extermination of the native caribou by 1955. Caribou were captured on nearby Coats Island in 1967 and airlifted to Southampton Island where 48 were released. The author studied the island vegetation in 1970 to 1972, as to forage quality and quantity in order to evaluate the carrying capacity of the range to support barren- ground caribou. By means of aerial photographs he divided the island into land-form types on the basis of moisture regimes and physiographic features. Parker follows F. A. Clement’s classical definition of plant formation as the climatic climax vegetation. Plant associations were considered the lowest unit for detailed description. The various plant associations were studied by the line-transect and plot method. In this case the line transects were 70 m long with fifteen l-m? plots selected at 5-m intervals. Data were collected on species frequency, cover and weight of the aboveground parts of the plants. Samples of lichens were analyzed for total nitrogen, caloric energy, and for phosphorus, calcium, magnesium, potassium, and Mammalogy By Harvey L. Gunderson. 1976. McGraw-Hill, New York. 483 pp., illus. $19.80. Harvey Gunderson of the University of Nebraska should be commended on writing a stimulating text on mammals. Unlike several recent books on mam- malogy, such as Cockrum’s /ntroduction to Mam- malogy and Vaughan’s Mammalogy, it makes no attempt to include extensive descriptions of the many living kinds of mammals; such information is valu- able, but it can be obtained in other sources, such as Walker’s Mammals of the World. Rather Gunderson concentrates on bringing together an interesting assortment of information from varied primary sources. The most innovative of Gunderson’s 16 chapters is that on the history of mammalogy as a science, beginning with its roots in European natural history and continuing with discussions on natural history in early America, the discoveries of explorers and fur traders, and finally the emergence of mammalogyasa discrete discipline. There is a useful section on collections of mammal specimens in United States museums and universities, on mammalogical work sodium in the laboratory. It was found that most herbaceous forage was produced in the limestone Hudson Bay lowlands while the most productive region for lichens was the Precambrian Plateau along the northeast coast. The greatest standing crop of lichens was approximately 1000 kg dry weight per hectare but the nutritional value of the lichens was low. It was concluded, however, that the quality of forage on the island was not a problem and an optimum caribou population for the island would be about 40 000 animals. The population was forecast to reach 1000 animals by 1980; afterwards, with the cooperation of the local Inuit, a modest harvest could be sustained. The Coats Island caribou were found to be exceedingly fat and the heaviest specimens for barren- ground caribou in Canada were recorded. Several males exceeded 180 kg in weight. The Coats Island population in March 1970 was approaching 2000 animals. A. W. F. BANFIELD Institute of Urban and Environmental Studies, Brock University, St. Catharines, Ontario L2S 3A1 done by American federal and state government agencies, and on journals devoted to research on mammals. Other sections of particular interest in this book are those on taxonomy and on physiology and behavior. Gunderson has used recent references on which to base his comments, so that his discussions are up to date, and often accompanied by well chosen figures and good photographs. It is understandable that in a large work of this scope some inconsistencies, repetitions, and errors will slip in; the use of both Lontra and Lutra for the otter; the statement that the camel’s temperature varies between 34° and 41°C repeated three times (temperatures are given in degrees Celsius, lengths and weights in both metric and non-metric units); and the use of the generic name Zebra. One must also contest Gunderson’s statement about Sir John Franklin’s last expedition in search of the northwest passage, “the earlier disappearance of whose members remains a complete mystery to this day” (p. 16). In fact Captain M’Clintock in his book The Voyage of the ‘Fox’ in the Arctic Seas (1859) described how the fate 204 of Franklin was discovered. (This book was reprinted in 1972 by Hurtig Publishers of Edmonton.) One small detail that bothered me was the use of a mammal’s scientific name on every possible occasion, especially such contentious ones as for the moose (Alces americana) and for the wolverine (Gulo luscus). The word pronghorn is followed by Antilo- capra americana over 20 times in the text. Surely in the interests of saving space, a list equating the BOTANY THE CANADIAN FIELD-NATURALIST Vol. 91 common and scientific names of a species could have been appended at the end of the book. Overall, this work is a fine one, up-dating previous texts on mammalogy. I am sure it will prove useful to future classes studying this subject. ANNE INNIS DAGG Box 747, Waterloo, Ontario N2J 4C2)5 Common Weeds of Canada/Les Mauvaises Herbes communes du Canada By Gerald A. Mulligan. 1976. McClelland and Stewart, Toronto (in association with /nformation Canada and The Department of Agriculture). 140 pp. Paper $4.95. The need has long been felt for a reasonably priced set of colored pictures of weeds. This completely bilingual book is definitely a step in the right direction. It contains 117 colored plates depicting 117 weeds of regional or widespread occurrence across Canada. Ten illustrations were reproduced from water-color drawings by the late Norman Criddle, first published in 1906 in Farm Weeds of Canada, now long out of print. The rest are from the author’s own colored photographs. Nearly all of the illustrations are about 3//2 X 5 inches (9 X 12 cm) and occupy the lower half of one page with the names and text above. Full-page illustrations are used for poison-ivy and common ragweed, the two weeds that figure most importantly as causes of human misery and dis- comfort in Canada. Most of the photographs are of a general view of one plant or a group of plants in natural setting. This has virtue in helping the reader form the valuable mental association of where a plant grows, together with what it looks like. But this also has the disadvantage of the plant being submerged into the background where the two are not in sharp contrast, either by color or by focus, with the result that some diagnostic features necessary for identifi- cation cannot be distinguished. Unfortunately, this happened in a number of the photographs, and their usefulness is less than might have been. Accompanying each illustration are the common and botanical names of the weed, the name of the family to which it belongs, and a paragraph including its life duration (annual, perennial), how it spreads, how high its stem usually grows, whether it is native in Canada or introduced, the provinces where it occurs, the habitats in which it is usually found, and the time of year when it flowers. The reader who seeks a more complete description of each weed is, in the Intro- duction, referred to the revised edition of Weeds of Canada. (This is an excellent book, published by Information Canada in 1970 with Mr. Mulligan as senior author, containing detailed descriptions with very accurate black-and-white line drawings.) Never- theless, had even a few of the more significant distinguishing characteristics of each weed been included in this new book, its value would have been much enhanced. This is especially true for those weeds whose features are not readily visible in the photo- graphs. Good quality paper was used and a nearly natural balance of color was achieved in most plates. Ina few, however, such as sulphur cinquefoil, the yellow was printed too lightly so that the petals turned out nearly white, and in others, especially the Mint and Composite families, the pinks and mauves lack the brilliance normaily associated with their flowers. Only one typographical error was noted: the second ¢ was missing from Bromus tectorum in the French column although it was spelled correctly in the adjacent English column. Attractively bound ina varnished black paper cover with an eye-catching group of 5 colored pictures on the front, this book will appeal to those who want a decorative volume as well as those who want help with identification of weeds. Being of pocketbook size, 5'/s X 8 inches (13 X 20 cm), it can be easily carried in the field, and, being completely bilingual, it should find ready acceptance in every province of Canada. JACK F. ALEX Department of Environmental Biology, Guelph, Guelph, Ontario NIG 2WI1 University of 1977 BOOK REVIEWS 205 Index to Plant Distribution Maps in North American Periodicals through 1972 Compiled by W. Louis Phillips and Ronald L. Stuckey. 1976. G.K. Hall and Co., Boston. xxxvil + 686 pp. U.S. $121. Plant distribution maps are invaluable resources for biogeographers and biosystematists, providing in- sights into evolutionary histories and barriers to interbreeding. For naturalists and conservationists, they indicate which species can be expected to occur within areas of interest and which may be locally noteworthy or rare. Taxonomists and naturalists generally will welcome the publication of this index. It will be useful not only for the specific purpose of locating maps but also as a guide to literature on the respective taxa. There have been numerous occasions in my own research and in dealing with requests for information when such an index would have been most helpful. The 268 journal titles listed (some merely name changes) represent a prodigious effort in searching as many North American periodicals carrying botanical articles as possible, including not only the well-known taxonomic journals but also many state academy journals, amateur naturalists’ magazines, and short- lived private publications. Thirty Canadian titles are listed. Because only journals published in Canada and the United States are covered, some maps of plant distribution in other parts of the world are indexed, and maps of North American taxa published elsewhere are not. A list of books containing plant distribution maps appears as a supplement. The 28772 entries appear alphabetically by scientific names, adhering to those used by the authors of the cited papers. Each entry includes the journal title, volume, page number, and year of publication, the type of map (e.g., dot or shaded), the geographic area covered, and the author’s name. Maps of both vascular and lower plants are indexed. This index is printed on good-quality paper, and is sturdily bound. Among books reproduced by offset from typed cards, it is extraordinarily neat and attractive. Its price will deter most individuals from purchasing this index, but it should be acquired by libraries wherever research in plant systematics and biogeography is conducted. JAMES S. PRINGLE Royal Botanical Gardens, Box 399, Hamilton, Ontario L8N 3H8 A Dictionary of Useful and Everyday Plants and Their Common Names By F. N. Howes. 1974. Cambridge University Press, New York. 289 pp. $12.95. Anyone who has ever used A Dictionary of the Flowering Plants and Ferns by J.C. Willis, 6th edition, 1931, will want to have a copy of this new book by F. N. Howes. When the already voluminous 1931 Dictionary was revised in 1966 so much new technical information on generic and family names had to be included that information on common names of piants and plant products had to be omitted. This omission was sorely felt by scientists and by users alike. F. N. Howes, then Keeper of the Museum of the Royal Botanical Gardens in Kew, England, under- took to fill that gap. He updated the material left out of recent editions by Willis and added so much new information that this volume now contains more than twice as many entries as before. Each entry is the English name ofa plant, part of a plant, a plant product, or the name of some person or condition associated with plants, or rarely, the botanical name of a plant. Entries are arranged in alphabetical order and have been accumulated from all over the world where English is or has been used or spoken. Each is accompanied by a brief but usually adequate description or definition plus the genus and species or just the genus of the plants involved. Where the same name refers to different products or species in different parts of the world, the country or region is indicated for each (e.g., Aust. for Australia, SE U.S. for southeastern United States of America, N. Am. for North America). There is a reasonable amount of cross-referencing as well. In many instances one common name merely refers to a second common name at which is found the definition and botanical name. Compound names, consisting of two or more words, are usually entered alphabetically by the first word only, e.g., “Thyme, common or garden Thymus vulgare” is defined under the “T’s” but “water thyme” (Anacharis, Elodea) is found only under “W”; “Ash pumpkin” appears only under “A,” not under “P.”- Fortunately, however, many entries are included under both letters, although sometimes incompletely so, such as “Prickly ash” which, under “A” refers to “Zanthoxylum spp.; Aralia spinosa:” but under “P” refers only to “Zanthoxylum spp.” Coverage tends to be better for plants of the United Kingdom, Africa, Australia, and the USA than of Canada or the Indian subcontinent. Although many Canadian native plants, cultivated introductions and introduced weeds are included, this seems to be largely 206 as a consequence of their occurrence in other parts of the world. For plants whose Canadian common name is the same as in the Old World there is no problem; but where different common names are used in the two regions for the same plant, it is frequently entered only under the Old World name. Or, where a particular common name applies to different plants in the two regions, the Old World definition is always given, but frequently there is no mention of the Canadian or North American usage. No large compilation like this can be entirely free of errors. Obvious typographical errors are virtually non-existent. Spelling errors are few and far between — an amazing feat when one considers the variety of A Panorama of Canadian Forests By Albert Potvin. 1975. Canadian Forestry Service, De- partment of Forestry, Ottawa, Canada. 254 pp., il- lustrated in color. Available from Information Canada, Ottawa. Canada $13.50, other countries $16.20. In this volume Albert Potvin has woven a brief outline of the forests of Canada about an absolutely striking series of color photographs, taken mostly by himself, from Newfoundland to British Columbia. The development of our forests is traced after glaciation to the eight forest regions recognized in Canada today. These regions are illustrated, both in close-up and from a distance. The role of fire and the elements and indeed the pressures of man are presented, together with a discussion of forest com- munities, tree distribution, and the inhabitants of the forest. The final chapter is a discussion of the future of Canadian Wildflowers By Mary Ferguson and Richard M. Saunders. 1976. Van Nostrand Reinhold, Toronto. 169 pp. $19.95. This is a delightful collection of beautiful color photographs of flowers that may be found in various parts of Canada, east, west, north, and south and from fields, forests, mountains, and arctic tundra. Many have previously been published in desk-calendar form as Canadian Wildflowers 1975, Canadian Wild- flowers 1976, and Canadian Wildflowers 1977 by Van Nostrand Reinhold, all authored by Mary Ferguson. The photographs were taken mostly by Mary Ferguson, but 15 other photographers have made contributions. Most of the pictures show only the flowers (or in some cases fruits) and not the whole plant, thus allowing for great depth of focus within the limited field. The colors are remarkably true. The THE CANADIAN FIELD-NATURALIST Vol. 91 sources from which this material was assembled: for example, Acacia cyanophylla was named after the Australian town, Cootamundra, but the spelling given here is Cootaminda; and the well-known Gray’s Manual of Botany, eighth edition, was authored in 1950 by M. L. Fernald, not by A. Gray. Nevertheless, this Dictionary is a most valuable compendium of information about “Useful and Everyday Plants” to Canadian users as well as to others throughout the world. J. F. ALEX Department of Environmental Biology, University of Guelph, Guelph, Ontario NIG 2W1 our forests, one of our most important possessions and a legacy to future generations. As a livingroom volume, A Panorama of Canadian Forests will receive much interest as viewers enthuse over the delightful pictures. It will, however, also be a pleasure for quiet comtemplation. The young student may too have his appetite whetted to develop his interest in our forested lands. To the stranger to Canada, it may foster a desire to visit this country to see it first hand. Ce livre est disponible aussi en frangais sous le titre Panorama des foréts du Canada. WILLIAM J. CODY Biosystematics Research Institute, Canada Department of Agriculture, Ottawa, Ontario KIA 0C6 ‘Canadian’ may be a slight misnomer, however, because there is a sprinkling of photogenic introduced species such as Purple Loosestrife, Bull Thistle, and Teasel. Each of the 144 species pictured is accompanied by an interestingly written description of the plant, its habitat and distribution, and items of particular interest concerning it. This is a “coffee table” book, which will certainly instil an interest in the delight of closely observing the flowers around us, in all those who turn its pages. WILLIAM J. CODY Biosystematics Research Institute, Canada Department of Agriculture, Ottawa, Ontario KIA 0C6 LOM, Rocky Mountain Flora By William A. Weber. 1976. Colorado Associated Univer- sity Press, Boulder, Colorado. 479 pp. $12.50. This is an excursion flora which any interested naturalist would find invaluable on a trip through Colorado, southern Wyoming, and northern New Mexico. Its popularity is evidenced by the fact that it is now in its fifth edition since first being published in 1953 as “Handbook of Plants of the Colorado Front Range.” The third and fourth editions were reviewed in this journal in 1968, Volume 82: 61; and 1973, Volume 87: 194. There are many changes in this edition: families are introduced with a paragraph of interesting and very readable information; many of the keys have been rewritten and species recently found in the region have been added so that about 1600 species, over half the Guide to the Vascular Flora of Illinois By Robert H. Mohlenbrock. 1975. Southern Illinois Uni- versity Press, Carbondale and Edwardsville. 494 pp. $7.95 (paperbound), 12.95 (clothbound). In 1967 Robert Mohlenbrock published the first volume of //lustrated Flora of Illinois: Ferns. This was followed by Flowering Rush to Rushes in 1970; Lilies to Orchids in 1970; Grasses: Bromus to Paspalum in 1972; and Grasses: Panicum to Danthonia in 1973; and Sedges: Cyperus to Scleria in 1976. It is obvious, however, that this series will take many years to complete at the present rate of publication. In the Guide to the Vascular Flora of Illinois, Mr. Mohlenbrock has taken the keys from the published volumes of the illustrated flora and those as yet unpublished, and consolidated them. To these he has added a key to families, brief descriptions of 14 physiographic regions under the headings of glacial history, bedrock, topography, soils, plant communi- ENVIRONMENT Superior: The Haunted Shore By Bruce Littlejohn and Wayland Drew. 1975. Gage Publishing Ltd., Toronto. 176 pp. $35.00 According to the authors, Superior: the Haunted Shore has two themes: “The first is the co-existence of power and fragility... ,” “The second is the humbling BOOK REVIEWS 207 flora of Colorado, are treated; observations on the ecology of some species have been adjusted as a result of field experience; new line drawings have been added; and the nomenclature has been updated. In this new edition the author laments the changes which have taken place throughout the region through the ravages and other results of urbanization. He has, however, provided a volume which has kindled an interest in the flora and through it may stay at least a part of the desecration of the country which has taken place since he began his studies of the flora of the Front Range. WILLIAM J. CODY Biosystematics Research Institute, Canada Department of Agriculture, Ottawa, Ontario KIA 0C6 ties, aquatic habitats, and distinctive fauna, indexes to common and scientific names, a glossary. Common names are given when known, occasionally very brief descriptions, notes on habitat, and when rare, locali- ties are given for individual species. The result is a handy little book which can readily be carried in the field. It will be most useful to students and naturalists who are interested in studying the flora of Illinois and adjacent regions. In Canada the book could be used to key out plant species in the southern parts of Ontario, for which unfortunately there is as yet no flora published, but the users in this province will of course find many species in it which do not occur in Ontario. WILLIAM J. CODY Biosystematics Research Institute, Canada Department of Agriculture, Ottawa, Ontario KIA 0C6 insignificance of the human record as the rocks of Lake Superior reveal it... .” It is the unique treatment of these two themes, however, which is the essence of this book. Each of the authors accepted one theme. “United in essentials, we therefore chose to work individually 208 during the writing and final selection of photographs, rather than striving for an integration which could easily become contrived and artificial.” The “power and fragility” theme is revealed through the excellent photographic efforts of Bruce Littlejohn. The book’s second theme, “the human record” is reported in the written words of Wayland Drew. The 98 color photographs selected by Bruce Littlejohn for inclusion are all of excellent photo- graphic quality. All portray the natural features of the Superior shore. All are of a high artistic merit. None portray people or the human impact and/ or existence on the Superior shore. If one is to find fault with the photography it would not be with the photographic skill of Littlejohn. One might suggest that there are too many similar-type shots included in the collection, such as waves crashing onto rocks. Shots I feel are missing are those from the numerous high lookouts along the north shore which show the vast magnitude of the lake and The Economy of Nature By Robert E. Ricklefs. 1976. Chiron Press, Portland, Oregon. 455 pp., illus. $13.95. It was a pleasure to examine Robert Ricklefs’ textbook in basic ecology. The book has been divided into nineteen chapters. The core idea of form, function, and factors is well established. The inter- dependence of the physical and biological realm was firmly believed to be the basis of the ecosystem concept. Examples were cited profusely throughout to present action, coaction, and interaction between organisms and against physical environment. The cycles of water, oxygen, carbon, nitrogen, and phosphorus were exhaustively dealt with. The book covers many minute details of every ecological phenomenon inevitably needed by a beginner. To my surprise, the topics like photoperiodism and circadian rhythms were omitted. This is not a drawback but they would have added interesting information to the total. Raunkiaer’s Life forms begin on page 73, but the Nature and Urban Man Edited by Gerald B. McKeating. 1975. Canadian Nature Federation, Special Publication Number 4. Ottawa, Canada. 184 pp. $4. This is a symposium volume comprised of nineteen papers. The stress is on the complex relationship THE CANADIAN FIELD-NATURALIST Vol. 91 the cold white-blue color of the water. Wayland Drew’s text is primarily historic, recount- ing legends and events, and describing people of times long past. Every so often reference is made to a Lake Superior site in a more current context, but this is a minor portion of the text. Whereas the Littlejohn photographs completely exclude people, the text of Drew is all of people. The writing is easy and enjoyable reading with the only noticeable flaw being the unfinished sentence on page 54. If this book has a fault it would be that only the north Canadian shore is treated. No consideration is given to the American Superior shore. Granted, the book’s present content makes it purely Canadian; however, to those who enjoy the world’s natural resources, nature’s treasures do not adhere to political boundaries. PETER CROSKERY Ontario Ministry of Natural Resources, Ignace, Ontario index shows page 75; this is a minor error. A table of conversion factors for area, length, time, mass, volume, velocity, energy, and power is included. A glossary giving all the definitions and meanings of ecological terms is added. At the end is a list of selected readings and text references for each of the chapters dealt with. These are quite useful. At the same time, had there been a listing ofall references (of Oosting, Phillips, Brown) about field techniques, it would have been very beneficial to the budding ecologist. On viewing all the above facts, I consider the book is quite readable, well composed, and balanced in subject matter, and is worth reading by anyone who is interested in ecology particularly, rather than by a student taking a course in a university. C. R. CHEVENDRA Science Librarian, University of Western Ontario, London, Ontario N6A 3K7 between man and the natural world. The purpose is to show some of the work undertaken and to identify some possible solutions. The contents are divided into five sections: overpopulation, capability of nature to survive in urban areas, the work undertaken by naturalists today, need for education and new approaches to planning and programs, and the 1977 approach by municipal and provincial authorities in Ontario. Specific examples were taken and discussed. Any conservation program should accommodate legitimately proper water management, recreational facilities, and wildlife management. A good sug- gestion was made, that is, the appointment of ecologists as staff members in professional schools of Law, Medicine, Architecture, Planning, and En- gineering to broaden the professional orientation in environmental issues and/or field ecology at second, third, and graduate level. A registry of professional ecologists is maintained in New Zealand and an appeal is made to Canadian authorities to follow it up for keeping a standard of environmental quality. The role of each professional in planning, policy-making, and program-developing is discussed, for example, OTHER Book REVIEWS 209 the role of an engineer, ecologist, public repre- sentative, biologist, etc. On the whole, the series of papers gives an insight into the various problems that exist in Ontario. Though all appear in tidbits, a theme is maintained consistently so that one can grasp the hazards of urbanization. The pros and cons of every little ecological problem under study were dealt with. I hope that the Canadian Nature Federation will continue its efforts with the same vigor in near future to bring out the degree and frequency of abuse during the battle between man and nature. C. R. CHEVENDRA Science Librarian, University of Western Ontario, London, Ontario N6A 3K7 Marine Sediment Transport and Environmental Management Edited by Daniel Jean Stanley and Donald J. P. Swift. 1976. Wiley, New York. xv + 602 pp. $35. This book by some twenty-three leading con- tributors is an outgrowth of lectures presented as a short course, ‘The New Concepts of Continental Margin Sedimentation, II,’ sponsored by the Ameri- can Geological Institute. Its target audience is ocean scientists, Ocean engineers, and the environmental managers representing the public who “must reconcile the competing demands made upon the (continental) shelf.” I remark that this latter description seems more appropriate to maximal utilization than to manage- ment in the broad sense. This book reflects a trend in the field toward a more holistic, dynamic approach. It is more holistic in that physical oceanography, chemical, and _ biological concerns are treated together with the sedimentology with some interpenetration of ideas. Dynamical oceanography occupies five early chapters. Sediment transport, both in bed load and in suspension, is discussed in equal detail. Patterns of sedimentation on beaches, at coastal inlets, on rocky coasts, at the continental shelf break (outer margin), in submarine canyons are similarly presented. The suite of six chapters on ‘Sedimentation and Environmental Management’ discusses sedimenta- tion with respect to beach and harbor engineering, structures, ocean mining, and ocean dumping. If one accepts the premise that it is man’s activities rather than the environment that is to be managed, then the words, ‘environmental management’ in this suite of chapters and in the book’s title would be better replaced by ‘environmental utilization.’ This volume is many textbooks in one. The marine practitioner can quickly choose for himself a set of chapters which will provide him a course of background reading addressing his current need. Some observations, offered in the Epilogue, help to place this work and this field of endeavor in perspective. Continental shelf sedimentology has been motivated largely by searches for petroleum and minerals. Studies of shore processes have been motivated by the requirements of coastal engineering. The field bespeaks bigness — in geographical extent, in expense, in the scale of man’s interventions, and in the epilogue writer’s projections for its future. There are, however, signs in this text and in the field that the natural limits are beginning to be perceived. Studies of the effects of dumped spoils on shelf waters and sediments are being actively pursued. The role of sediments in supplying nutrients for shelf fisheries is beginning to be appreciated. RONALD H. LOUCKS R. H. Loucks Oceanology Ltd., 24 Clayton Park Drive, Halifax, Nova Scotia B3M 1L3 210 THE CANADIAN FIELD-NATURALIST Vol. 91 NEW TITLES Zoology Amphibians and reptiles in Alaska, the Yukon and North- west Territories. 1976. By Robert Parker Hodge. Alaska Northwest (Canadian distributer, Hurtig, Edmonton). 92 pp., illus. Paper $4.75. + Annotated checklist of the birds of Ontario. 1976. ByR. D. James, P. L. McLaren, and J. C. Barlow. Royal Ontario Museum, Toronto. 75 pp. $2.50. + Biocides and birds. A survey of the effects of biocides on birds. 1976. By R. E. Rogers. Alberta Environmental Con- servation Authority, Edmonton. vii + 90 pp. Paper free. Biology of opisthobranch molluscs. Volume |. 1976. By T. E. Thompson. Ray Society, London. vi + 208 pp., illus. £15. { The biochemistry of blue, snow and Ross’ geese. 1976. By Harold C. Hanson and Robert L. Jones. South Illinois University Press, Carbondale. xvii + 281 pp., illus. $15. The birds of Alberta. With their ranges in Saskatchewan and Manitoba. 1976. By W. Ray Salt and Jim R. Salt. Hurtig, Edmonton. 2nd edition. 512 pp., illus. $10. + The bluebird. How you can help its fight for survival. 1976. By Lawrence Zeleny. Audubon Naturalist Library Book. Indiana University Press, Bloomington. xix + 170 pp., illus. $7.95. The broken archipelago. Cape Cod and the islands, amphi- bians and reptiles. 1976. By James D. Lazell. Photographs by Martin C. Michener. Quadrangle (New York Times), New York. xi + 260 pp. $12.50. Butterflies of West Malasia and Singapore. 1975. By W. A. Flemming. Classey, Faringdon, England. Two volumes, illus. x + 94 pp. + plates, and x + 64 pp. + plates. $47. + Crows of the world. An authoritative guide to all 116 species of the crow family, including jays, magpies, rooks, and the raven. 1976. By Derek Goodwin. Illustrated by Robert Gillmor. Cornell University Press, Ithaca, N.Y.vi+ 354 pp. $28.50. {+ A guide to birdwatching in Mallorca. 1976. By Eddie Watkinson. Available from M. Philbrick, Box 83, Vashon, Washington. 56 pp., illus. $3.90. Insect clocks. 1976. By D.S. Saunders. Pergamon, New York. International Series in Pure and Applied Biology. Volume 54. viii + 280 pp., illus. $18.50. Insect flight. 1976. Edited by R. C. Rainey. Symposium of the Royal Entomological Society of London Number 7. Halsted (Wiley), xii + 288 pp., illus. $47.50. Island biology illustrated by the land birds of Jamaica. 1976. By David Lack. University of California Press, Berkeley. xviii + 446 pp., illus. $25. Marine insects. 1976. Edited by Lanna Cheng. Elsevier, New York. xi + 582 pp., illus. $62.50. Marine mussels. Their ecology and physiology. 1976. Edited by B. L. Bayne. Cambridge University Press, New York. xvii + 506 pp., illus. $49.50. Moths of Southern Africa. Descriptions and colour illustra- tions of 1183 species. 1975. By E. C. G. Pinhey. Tafelberg, Cape Town. x11 + 274 pp. + plates. $35.95. A natural history of zebras. 1976. By Dorcas MacClintock. Pictures by Ugo Mochi. Scribner, New York. x + 134 pp. $7.95. Report of the committee on the killing of wild birds for scientific and educational purposes. 1976. By the Pan- American Society for the Protection of Birds. Available from Dr. J. B. Tatum, Department of Physics, University of Victoria, British Columbia. 14 pp. $1. Social behavior in vertebrates. 1976. By Anthony Payne. Heinemann, London. vi+ 146 pp., illus. $6.75. Sphecid wasps of the world. A generic revision. By R. M. Bohart and A. S. Menke, in collaboration with H. S. Court, F. D. Parker, E. Grissell, and D. P. Levin. University of California Press, Berkeley. x + 696 pp., illus. $42.50. Where the grizzly walks. 1977. By Bill Schneider. Moun- tain Press, Missoula, Montana. $8.95. Wintering bald eagle. 1976. By Donald A. Spencer. National Agricultural Chemicals Association, Washington. ix + 170 pp. Wolf...kill! The wilderness called Shunka. 1976. By Marika Lumi. Van Nostrand Reinhold, Scarborough, Ontario. 195 pp. $9.95. Botany {+ Canadian forestry. The view beyond the trees. 1976. By Charles R. Stanton. MacMillan, Toronto. 70 pp., illus. Paper 5.95. Canadian wildflowers. 1976. By Mary Ferguson and Richard Saunders. Van Nostrand Reinhold, Scarborough, Ontario. 192 pp., illus. $19.95. Common mosses of the Pacific coast. 1975. By Marion P. Harthill and Irene O’Connor. Naturegraph, Healdsburg, California.115 pp. Evolution of crop plants. 1976. Edited by N. W. Sim- monds. Longman, New York. 339 pp. $44.37. 1977 A field guide to Pacific States wildflowers. Field marks of species found in Washington, Oregon, California and adjacent areas. A visual approach arranged by color, form, and detail. 1976. By Theodore F. Niehaus. Illustrations by Charles L. Ripper. The Peterson Field Guide Series, Number 22. Houghton Mifflin, Boston. xxxii + 432 pp. $10.95. Flora of Guatemala. 1976. By Dorothy L. Nash and Louis O. Williams. Field Museum of Natural History, Chicago. Fieldiana: Botany, Volume 24, part 12. x + 604 pp., illus. Paper $18. Nova Scotia boletes. 1976. By Darryl W. Grund and Kenneth A. Harrison. Bibliotheca Mycologica 47. Cramer, Vaduz, Germany. 283 pp., illus. * Ontario weeds. Descriptions, illustrations and keys to their identification. 1976. By J. F. Alex and C.M. Switzer. Ontario Ministry of Agriculture and Food, Toronto. 200 pp., illus + plates. $2.50. Origin and evolution of angiosperms. 1976. Edited by Charles B. Beck. Columbia University Press, New York. 341 pp. $17.50. Seeds and fruits of North American Papaveraceae. 1976. By Charles R. Gunn and Margaret J. Seldin. Agricultural Research Service Technical Bulletin Number 1517. United States Department of Agriculture, Washington. 96 pp. Wild edible plants of the western United States. 1975. By Donald R. Kirk. Naturegraph, Healdsburg, California. 307 pp. Wild harvest. An outdoorsman’s guide to edible wild plants in North America. 1975. By Alyson Hart Knapp. Illus- trated by E. B. Sanders. Pagurian, Toronto. 190 pp. $8.95. Woody plants of oak openings. 1976. By Bill Easterly. Illustrated by Joy Marburger. Available from author, 506 Harvest Lane, Bowling Green, Ohio. $6.50. Environment Biogeography and ecology in the Canary Islands. 1976. Edited by G.G. Kunkel. Monographiae Biologicae, Volume 30. Junk, The Hague. xvi + 512 pp., illus + plates. Dfl. 160. Citizens and the environment. Case studies in popular action. 1976. By Lynton K. Caldwell, Lynton R. Hayes, and Isabel M. MacWhirter. Indiana University Press, Bloomington, Indiana. xxx + 450 pp. $17.50. * The ecology of the seas. 1976. Edited by D. H. Cushing and J. J. Walsh. Saunders, Philadelphia. 467 pp., illus. $19.50. BOOK REVIEWS 4 211 Environmental knowing. Theories, research and methods. 1976. Edited by Gary T. Moore and Reginald G. Golledge. Dowden, Hutchinson and Ross, Stroudsburg, Pennsylvania. Xxil + 442 pp., illus. $25. Insights into environmental education. 1976. By George C. Martin and Keith Wheeler. Oliver and Boyd (Longman), New York. xii + 200 pp. $11.50. Proceedings of an international conference on marine parks and reserves. Tokyo, May, 1975. 1976. IUCN Publications New Series, Number 37. International Union for Conserva- tion of Nature and Natural Resources, Morges, Switzer- land. 132 pp. Paper $6. Resource conservation glossary. 1976. By Soil Conserva- tion Society of America, Ankeny, Iowa. 63 pp. $5. Sea life of the Pacific northwest. 1976. By Stefani Hewlett and K. Gilbey Hewlett. Illustrations by Greg Davies. McGraw-Hill, New York. 176 pp. $14.95. Miscellaneous Concepts of species. 1976. Edited by C. N. Slobodchikoff. Benchmark Papers in Systematic and Evolutionary Biology, 3. Dowden, Hutchinson and Ross, Stroudsburg, Penn- sylvania. 368 pp. $10. The conservation response. Strategies for the design and operation of energy-using systems. 1976. By Lloyd J. Dumas. Lexington (Heath), Lexington, Massachusetts. xvill + 290 pp. $14. Field photography. Beginning and advanced techniques. 1976. By Alfred A. Blaker. Freeman, San Francisco. (Canadian distributer Oxford University Press, Don Mills). xxi+ 451 pp. + 41 pp. field book, illus. $19.95. Richard Harrington’s Antarctic. 1976. By R. Harrington. Alaska Northwest. (Canadian distributer, Hurtig, Edmon- ton). 104 pp., illus. Paper $9.95. +The Shetland way of oil. Reactions of a small community to big business. 1976. Edited by John Button Thuleprint. Sandwick, Shetland. 134 pp., illus. £2.40. The world of rocks and minerals. 1976. By Anita Mason. Photographs by Eric Storey. Larouse, New York. xviii + 108 pp. Cloth $8.95, paper $4.95. The world of remote sensing bibliographic index. 1976. Compiled by Paul F. Krumpe. Tensor, Merrifield, Virginia. 600 pp. $18. + Assigned for review * Available for review The Ottawa Field-Naturalists’ Club Minutes of the Ninety-seventh Annual Business Meeting of The Ottawa Field-Naturalists’ Club The 97th Annual Business Meeting of The Ottawa Field-Naturalists’ Club was held in the auditorium of the National Research Council, Sussex Drive, Monday, 19 January 1976. The President, E. C. D. Todd, called the meeting to order at 8:10 p.m., witha quorum of 38 persons present (late arrivals brought this to 47 in all). After the agenda was approved on motion (by R. Taylor, 2nd I. Brodo), the Recording Secretary read the minutes of the 96th Annual Meeting. No errors or omissions were noted, and the minutes were approved on motion (by A. Erskine, 2nd C. Gruchy). The President called on C. Gruchy to introduce the amendment to the Constitution that was to be voted on; Gruchy explained that its purpose was to meet the legal requirements for recognition of the Club as a “charitable organization” which could issue tax-free receipts for donations received. This amendment, moved and seconded at the last annual meeting, was approved after brief discussion. R. Foxall, chairman of the Finance Committee, spoke on the financial status of the Club. He explained that, owing toa recent transfer of the Club’s funds from a trust company to a bank, and to late receipt of year-end statements from the bank, the Treasurer had been unable to complete compilation of the financial statement in time for this annual meeting. Since the Constitution calls for presentation of this statement at the annual meeting, a motion (by C. Gruchy, 2nd D. A. Smith), empowering Council to receive the financial report instead at the earliest possible opportunity, was approved. The statement will appear in The Canadian Field- Naturalist as usual, following acceptance by Council. There was a suggestion that the meeting be held later in January to allow more time for preparation of-the financial statement, and an enquiry about the loss of interest incurred by the transfer of invested funds between banking facilities. No estimate of such loss was available, but Foxall emphasized that the transfer had been necessary because of poor service by the former facility and rumours of its financial instability. E. Todd then introduced the annual report of Council, which is published in The Canadian Field- Naturalist, and commented on some of its highlights. These included re-organization following a study carried out by H. MacKenzie; a bequest of $500.00 from the estate of the late Rowley Frith; the choice offered to individuals of membership in the Club or of subscription to The Canadian Field-Naturalist with no membership privileges; the weekend excursion to the St. Lawrence estuary to see whales and seabirds; and the activities of the Conservation Committee, the Macoun Field Club, and the Centennial Planning Group. Its adoption was moved (by T. Mosquin, 2nd I. Sutherland) and passed. A. Reddoch, chairman of the Nominating Com- mittee, noted that the committee had followed the guidelines of the draft by-law on nominations in assembling a slate, as follows: President: Ewen Todd; Vice-President: Roger Foxall; Treasurer: Pamela Sims; Recording Secretary: Anthony Erskine; Corres- ponding Secretary: Patricia Narraway; additional members of Council: Elisabeth Beaubien, William Cody, Albert Dugal, David Gray, Diana Laubitz, Hue MacKenzie, Diane McClymont, Jo Ann Murray, Gerald Oyen, Roger Taylor, Stanley Teeple, Stan Van Zyll de Jong. A request for nominations published in Trail & Landscape \ed to one additional nomination from the general membership: Marshall Ney. On motion (by A. Reddoch, 2nd E. Dickson) all of these nominated were declared elected. R. Foxall expressed his approval of the fact that a nomination had been received from the membership at large, and hoped that more persons would be so nominated in future. L. C. Smith queried the status of past presidents on Council. The consensus was that one year’s continua- tion by an outgoing president helped continuity of operations, that a longer period was unnecessary for that purpose, and that no formal statement of this was required. The President introduced both old and new Council members then present, and thanked outgoing members of Council: J. Dafoe, C. Gruchy, E. Haber, D. Lafontaine, L. Padelford, A. Reddoch, J. Red- doch, A. Sheppard, F. Weekes, H. Williamson, and I. Brodo. He also thanked the auditors for 1975, G. D. Tippett and G. J. Wasteneys, and the refreshments organizer, Catherine O’Keefe. The auditors for 1976, G. J. Wasteneys and D. A. Potter, were approved on motion (by R. Foxall, 2nd C. Gruchy). Under new business, D. Gray asked for volunteers to help with various aspects of the program of the Macoun Field Club, especially in connection with excursions, summer trips, and the publication The Little Bear. E. Todd introduced T. Mosquin, Execu- tive Director of the Canadian Nature Federation, who was seeking volunteers to help with the organization’s conference in Ottawa in May 1976. G. Neville raised questions on financial procedures, and R. Foxall explained that with the increased size of the Club, the work of the Treasurer required as much or more time as several other positions for which honoraria are now granted; the Finance Committee will consider whether the workload should be split or some other procedure would be more appropriate. G. Neville also IND 1977 queried action on the Club’s centennial projects, with particular reference to the possibility of land acquisi- tion. R. Foxall noted that this idea had been con- sidered relatively low in feasibility by members of Council at the special meeting held to consider various proposals for centennial projects, but acknowledged that changed circumstances in future could make it more feasible. In further discussion, it was noted that other proposals for centennial projects dealt with specific items or events, whereas only the idea of acquiring land had been advanced. No particular tract had been proposed for consideration, and Council believed that any tract of a size sufficient to be ecologically viable or recreationally valuable would be so costly as to rule out the possibility of any other major project being undertaken concurrently. G. Neville noted that once donations to the Club were tax-free it would be possible to mount an appeal for funds for this purpose, without committing current funds for the purchase. Problems in land management experienced by other clubs holding land were also THE OTTAWA FIELD-NATURALISTS’ CLUB 213 mentioned; M. Stuart noted that a piece of land made available to the Club in the past had later been disposed of after it proved too difficult to maintain. Following a suggestion by T. Mosquin, a consensus was reached that a meeting of the Club be held, with representatives invited from the Federation of Ontario Naturalists’ nature reserves program and from other clubs holding land for nature purposes, to discuss the topic in depth. R. Foxall noted that no centennial proposal had yet been finally eliminated from consideration, but that a decision on major projects would be needed before the next annual meeting. H. MacKenzie noted that some projects are already being organized by Council, and such action should be reported in Trail & Landscape, without waiting for the next general meeting. The meeting was adjourned on motion (by I. Sutherland) at 10:00 p.m. A. J. ERSKINE, Recording Secretary Report of Council to The Ottawa Field-Naturalists’ Club In 1976, the Centennial Steering Group pointed out that a number of policy decisions were needed from Council so that the planning group was not placed in the position of determining Club policies for the next several years. The Council decisions were these: 1. Focus—both the local and national roles cur- rently assumed by the Club should be con- tinued. Rather more emphasis should be placed on the local role, with the national role per- formed mainly through The Canadian Field- Naturalist. 2. Membership—we should seek more active and involved members. This need not involve any major change in total numbers. 3. Participation—both the public and Club mem- bers, but especially the latter, should be able to participate in the Centennial. 4. Nature of observance—the Centennial should include both serious and fun activities. 5. Perspective—the balance is in favor of retro- spective topics but there is also strong support for looking ahead. With these broad guidelines, the Centennial Steering Group is endeavoring to plan a program of activities which will have “something for everyone.” This need for planning led to Council’s establishing an Executive Committee, which had been one of the recommendations of the study of Club policies and practices reported last year. Discussions in this committee should help better to focus subsequent consideration of a topic by Council. Reports of other committees follow, with names of chairmen in paren- theses. Finance Committee. Early in the year, the Finance Committee prepared the budget for 1976. The change in Club policy, whereby a member was provided the choice of either remaining a member or taking an individual subscription to The Canadian Field- Naturalist, resulted in a decrease in membership with more money going directly to the journal. Accord- ingly, the proportion of membership dues used to support CF-N was reduced from 50% to 40%. Con- sideration of the Club’s financial picture in 1976 led the Committee to recommend to Council that the fee schedule for 1978 be increased by $2.00. A bequest of $250 was received from the estate of Louise Gourlay, an active member of the Club for 24 years. After extended negotiations the Club has finally achieved official status as a “charitable organization”; receipts for income tax purposes can now be issued for donations to the Club. (R. Foxall) Membership Committee. Members had been offered the choice of becoming Subscribers to The Canadian Field- Naturalist without membership privileges in the Club, or continuing as Members of the Club, which entitled them to participate in Club activities, hold 214 office, vote, and receive Trail & Landscape as well as The Canadian Field-Naturalist. This choice subse- quently resulted in a large drop in membership for 1976, amounting to 36% among non-local members but only 5% of local members (see Table for the comparisons with 1975 figures). Membership appli- cation forms continue to be a source of information concerning members who volunteer their skills or specialized knowledge in assisting the Club. Thanks go to all members who volunteered their services. Increased postal rates, printing, and paper costs continue to rise, and this will affect future member- ship fees. Honorary Membership for W. Earl Godfrey was recommended, and approved by Council. (M. Ney) Publications Committee. Four issues of The Cana- dian Field-Naturalist, Volume 89(4), and Volume 90(1,2,3), totalling 538 pages, were published since the last report. The last issue, dedicated entirely to raptors, deserves special mention. Articles and notes published during the period numbered 103, including 41 on ornithology, 22 on mammalogy, 14 on botany, 8 on ichthyology, 5 each on herpetology and limnology, and 8 on other subjects. Also 49 book reviews were published. The number of manuscripts submitted to the Editor in 1976 totalled 147, about the same as in the preceding period. We are grateful for grants in support of the journal received from the National Research Council ($10,000) and the Canadian National Sportsmen’s Show ($750). Five issues of Trail & Landscape were published, comprising 140 pages, and covering a wide range of subjects and information of interest to the naturalist. The publi- cation of Trail & Landscape was supported by a $250 grant from the Canadian National Sportsmen’s Show. (S. Van Zyll de Jong) Excursions and Lectures Committee. During 1976 ten monthly meetings and 37 field trips were organ- ized. These latter focussed on the following subjects: birds (20), general (7), botany (5), insects (2), rocks and fossils (2), and amphibians (1). The evening meetings dealt with various natural history topics such as spring wildflowers, mineralogy, and herptiles, and included a panel discussion, organized by G. Neville, on the question of land acquisition. In addition, the Club’s annual dinner was held at the Talisman Inn on 20 April. The speakers were John and Janet Foster on “The Wild Regions of Canada.” Approximately 300 people attended. The Club also arranged and led a number of excursions for the Canadian Nature Federation meeting in Ottawa in late May 1976. (R. Taylor) THE CANADIAN FIELD-NATURALIST Vol. 91 Conservation Committee. Many communications were received seeking information, or transmitting it, or soliciting commentary on proposed plans. Not all those issues deserving comment received it, since time to study or respond was seldom available when needed. The potential activities of this committee are limited only by the time its members can find for it. A brief on the conceptual plan for Gatineau Park, prepared by Allan and Joyce Reddoch after a dis- cussion in committee, was presented at the public hearing in November. Other briefs were submitted on mineral aggregates extraction policy in Ontario and on recreation policy in Canada. Questionnaires or enquiries responded to included the Club’s uses of conservation areas designated in the Ottawa-Carleton Region Official Plan, values of the Ottawa Greenbelt, and natural areas in the southeast part of the City of Ottawa. Major planning issues on which no sub- mission was made included the Outaouais Region plan, the N.C.C. interpretation planning for the Greenbelt, Nepean Township’s plan for the Jock River corridor, and the City of Ottawa open space study. These mostly involved 50- to 100-page presen- tations, some making use of data previously made available by the Club, and superficial examination revealed few obvious weaknesses. (A. Erskine) Macoun Field Club Committee. This year members actively continued the Macoun Club’s program for young naturalists, including field trips, meetings, and the Seniors’ summer canoe trip. During the summer Len Marhue resigned as chairman, and Jerry Fitz- gerald (NMNS) and Arnet Sheppard (OFNC) took over as co-chairmen. The membership remained at about 90. Juniors and Intermediates had weekly meetings and monthly excursions, while the Seniors had both meetings and field trips weekly. The Seniors also helped other groups by conducting nature walks, planning and building nature trails, and surveying new areas. Reports on Club activities will appear in the 1976 “Little Bear.” Thanks are due to the National Museum of Natural Sciences (co-sponsor) and to parents of Club members who helped in various ways. (D. Gray) Education and Publicity Committee. This committee concentrated on providing trip leaders for outside groups, predominantly Cubs, Scouts, and Guides. Macoun Field Club seniors were often effective as leaders. A number of OFNC members braved the mud and sleet to set up telescopes for the two weekends of the Ottawa Duck Club’s open house. Owing to confusion about the date of judging at the Ottawa Region Science Fair, it was impossible to award 1977 individual prizes; instead a contribution was made to help the overall winner in natural science to meet travel expenses to the National Science Fair. (A. Sheppard) The support of the winter bird feeders on Moodie Drive (west end) and Davidson Road (east end) has been continued, and in the fall of 1976 a third feeder near Pink Road in Lucerne was added. A new birding newsletter named “The Shrike,” THE OTTAWA FIELD-NATURALISTS’ CLUB Als) produced by Club members but without Club financ- ing, appeared first in 1976. (S. Van Zyll de Jong) Thanks are extended to all persons who have served the Club in various ways during 1976. Compiled from committee reports and Council minutes by A. J. ERSKINE, Recording Secretary Membership of The Ottawa Field-Naturalists’ Club Canadian Foreign Category Local Other U.S.A. Other Totals 1975 1976 1975 1976 1975 1976 1975 1976 1975 1976 Individual 451 408 542 324 127 85 09 6 1129 823 Family 198 203 16 23 2 3 0 1 216 230 Sustaining 5 9 l 2 0 0 l 0 7 11 Life 6 7 ] ] l 2 2 10 14 Honorary 5 5 4 4 0 0 0 0 9 9 Totals (on 25 Nov./76) 665 632 564 357 130 89 12 9 1371 1087 Changes -33 207 -41 -3 284 Auditor’s Report To: Members of the Ottawa Field-Naturalists’ Club We have examined the balance sheet of The Ottawa Field-Naturalists’ Club as at December 31, 1976 and the related Income Statements for the year then ended. Our examination included a general review of the accounting procedures and such tests of the records and supporting vouchers as considered necessary in the circumstances. In our opinion these financial statements present fairly the financial position of the organizations as at December 31, 1976 and the results of their operations for the year then ended in accordance with generally accepted accounting principles. (Signed) Geoffrey Wasteneys D. A. Potter CGA January 28, 1977 216 THE CANADIAN FIELD-NATURALIST The Ottawa Field-Naturalists’ Club Balance Sheet as at December 31, 1976 Assets Current @asheinebank= OFEINi Cairne rice $ 2,758.79 (GAG ih yan <=(CJRIN, Sob oounnotoooooneoduocgcDoc 21,537.14 Billsirecenvalble siete a ree renessr cnet ale aires banc eeronnteaee Accnuedsintenestinecenvalbleminer een orien eae PirSoeNG! GOINGS“ oaccdconandecoboovcouccngc00c00 Fixed (at cost) Furniture, fixtures and equipment ................. Messsaccumulated depreciation) 4-4. eee Investments and securities CanadasSavinespBondsieeneeee neces cient Current liabilities Income received in advance ............-0.-e000%> ACCOUNTS Payable errs chateaus you cus cusvercbe) svete Equity of surplus names Jemunamy Il, IDVO udosavcsssaccodencosuceue Addemetencomextorthenyecal ae ace cee accra Vol. 91 $24,295.93 3,737.01 7,896.15 4,863.00 $40,792.09 529.50 390.70 138.80 10,700.00 $ 6,223.50 6,948.59 $13,172.09 26,747.73 11,711.07 38,458.80 $51,630.89 (Signed) Geoffrey Wasteneys, Auditor D. A. Potter, Auditor Pamela J. Sims, Treasurer 1977 THE OTTAWA FIELD-NATURALISTS’ CLUB PAW) The Ottawa Field Naturalists’ Club Statement of Profit and Loss— CF-N for the year ended December 31, 1976 Revenue Memibershipmncomen enn acccise ie sec icrelrecie: SUSHI INXS GoopodacoccuobouoodGdnhoboodue Grants—N.R.C. —Canadian Sportsmen Show ............... IRC = Sad cie testo cee obra cee coolio mrolaalcae recto siaees Rlatessandatabrsettings, Sssesnemocesceaeaci sce ace Extra pages and authors’ costs ...............-.-0- BB ACKSMUIMIDETS eve ac. sees jabs vcar'e sensi wits o dalla ays ca snieueh see enoeeuonese Special publications MMC SNES tT OTM Caesar agers ortiases cette tonsvoye a ie eeehiev susucvane a ets ee Ce a CC Py Less: cost of publications Volume 90 (1, 2, 3, 4) latesvandatalbssettinGs) = oecmssecine cre chests siecusiere ING RIMUNC OS tS racsesery ace, inate ciate apalls o: wcchastoone ears ave eters Gross profit on operations Less: operating expenses Bank charges and interest ...............-.-eeeeee W@incwlatronie stele doen eee sale Wreeoe leeds Office assistant AOS PAE Ciiactentace orsie satel scieiis/n to sstarelois Gis wieiinle. sicrausineniece¢ Printing and stationery Editing == Ontia Cts) aajs cisco ain cx « levctenslelas) ole ejeraieiieyereve —General expenses Honoraria Ca Net Income $ 4,296.13 10,775.30 $10,000.00 750.00 9,918.25 3,053.00 5,019.12 2,727.45 300.00 1,606.83 26,034.72 112.00 4,589.29 21.41 900.11 1,518.00 1,819.32 857.07 890.00 739.74 1,502.00 10,750.00 $48 446.08 30,736.01 17,710.07 82,247.65 $ 9,462.42 218 THE CANADIAN FIELD-NATURALIST Vol. 91 The Ottawa Field Naturalists’ Club Statement of Profit and Loss— O.F.N.C. for the year ended December 31, 1976 INetsincomefromiGReNi eas oe cerca ere aes as) er eee or Other income IMIG EIN} WNCOINS Gobo cooneccacadsassoon suede ono odeuoondound $6,451.20 Donations anderantsy 125 Som 3e Sees eal coaey vena cater syeromeve enlace sewsnsrane 515.94 Sal O11 CO MV Scene cases sree eee cee eee art alae epeeh ack Guna hens cpatouseher ame ae wenaeaecewe 8.00 Subseriptiomsy Dy ole fe ih. eh ees ene cceeele eiiede ve pees a couteaeals Shcasnoneaeret 173.75 Intenestaincomerandadividend Sierra eerie 660.10 Less: cost of publications JE Cee CoN KON WG COV eS Daa ais icnsancats teed eens uo Cnols eo conic ond camino era ob 2,079.15 (Gee UI ES Voy alten cae osetia incr icin cates ee ae carte aI cr Renna aoe Maan ce 120.80 Officevan dhe dito riallicaks cre seep ees eve. epee sich a esse ne omer ee esos acute 61.19 LOM OTe Raa toh ee ia tect (econ aca oy era valcre! even sucha eet S cal ars amaens Pata ues oR 440.00 Gross profit on operations ........5 0.0... c eee cee rete eee eee Less: operating expenses CouneilFexpenSesise capes aeeaxs sear aseie eatieed ues eer daa: taparovet bean, Se dovemntoe 964.79 iMtingsandystatlOnenyie eves: sisccuersieh 100) WHOYIUAIS = ge (10 Sd > SOO) RdYIUaIS AjeUIdIEW = 4 WUROIJIUTIS JOU = “s'U :sMO|[OJ se samu[Iqeqoud palier-OM | *sasnsuad Jo Jaquinu — N, -|z| = sonsiieas say ‘OZ SN JOyra uaym “OZ > 1461 PUB L961 YI0G 10} N Udy F) AoUTTyA\-UURW = oNSHeIS IS9], c01}S1} 3S 489], ‘saioads yora Jo sduljyais yse] pue ssly ay UI9MJ9q Spoliad Aep-¢ aAIsnjoul Jo Jaquinu = N z "€ OINBIy Wosy sanjeA 19q0j}9Q-Jaquiaidag pur isndny—Ain¢ ayy Jo adesaay (PEEL FS IZ (PE LIFSLL (p) 8S +87 (Lp F8TI (b)S0 #80 (p) OSI FS°L9 (bp) € 8E + 0'°86E (pb) L6€ £ 8°ZOl (p) PEL HSL (Pp) €6€ FE IP (yp) 911 $06! 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FHP (8) 81 #8 (LD 60 +10 (8I)7% FET 1L61 L961 potiod Aep-¢ tod snsuao Jad AN) “A'S F ueap julog duo] € auojsuiny Appny 8 Sul[apues ve urjund Le T9AO[d P2[194-A9ePlA salgads a0Rjiajul yovog LL Jodidpues [e10399q SL Jadidpues s,piieg €8 I2A0[q payewyediuas +6 Jadidpues pajewyediuas 96 Jadidpues iseoq 66 S3I[MO][IA asso] 66 SBI[MOTJAA 19}891H) 00! ‘dds 194y911IM0q saidads ood yovag Jadidpues pajyjods TOP I9A0[q Budi soisods Susan sa19eds 1L961 90 —"In¢ sjood yoroq uo % 1L61 PUL £96] U2EMIAQ SUT[OIOYS d11q axe] UID}SeOYyIIOU dy} pUe JUIOg BUOT 3 spiqoioys Jo souRepUNe 10qQ0}90 0} A[NE ay} UI UONBIeA—p ATaV 234 October in 1967 and in 1971. Since Page and Bradstreet (1968) have shown that marked Least and Semi-palmated Sandpipers may remain on Long Point for many days during migration, daily census data are not statistically indepen- dent for these and probably other species. To improve independence and minimize the skew- ness that consideration of 5-day peak counts would cause, the mean number of birds seen per count in each 5-day interval was used as the unit of observation in comparison of the 1967 and 1971 data. The sample size in each year was the inclusive number of 5-day intervals between the first and last sightings of a species. The Buffalo Ornithological Society counted shorebirds in two areas totalling approximately 32 km of northeastern Lake Erie shoreline in 1967 and 1971 (Seeber 1967; A.R. Clark, unpublished data) (Figure 1). Three counts were made in 1967 (22-23 July, 18-19 August, and 2-3 September) and four in 1971 (18 July, 8 and 22 August, and 5 September). The mean number of shorebirds per count is used to compare the abundance of the birds between the two years. For comparisons between years in both areas, Mann-Whitney U tests (Siegel 1956) are used. Variations in shorebird abundance in 1967 and 1971 are then compared between Long Point and northeastern Lake Erie. Nesting Species. Mean numbers of Killdeers at Long Point did not differ significantly between 1967 and 1971 (P>0.1). significantly fewer (P<0.01) Piping Plovers were seen in 1971 than in 1967. Snyder (1931) estimated 100 nesting pairs of Piping Plovers at Long Point in 1927 and 1928, and Hussell and Montgomerie (1968) about seven pairs between 1961 and 1965. By 1971, only four adult Piping Plovers were present and to our knowledge only one young was fledged. Significantly more (P<0.01) Spot- ted Sandpipers were seen in 1971 than in 1967. Beach Pool Species. All eight species that fed primarily at beach pools during the southbound migration in 1967 were less abundant in 1971 (Table 4). This decline was highly significant (P<0.01) for Greater Yellowlegs, Least Sand- pipers, and dowitchers, significant (P<0.05) for Semipalmated Plovers, Baird’s and Semipal- mated Sandpipers, marginally significant (0.1>P>0.05) for Lesser Yellowlegs, and not THE CANADIAN FIELD-NATURALIST Vol. 91 significant (P>0.1) for Pectoral Sandpipers. Considering all eight species combined, only 23% as many individuals were seen on an average census in 1971 as in 1967. In contrast, 81% as many were seen in 1971 as in 1967 along the northeastern shoreline of Lake Erie. Beach Interface Species. For species that fed primarily along the beach-lake interface at Long Point, no significant changes were found in autumn numbers of Black-bellied Plovers, Ruddy Turnstones, or Dunlins (Table 4). The increase in the numbers of Sanderlings between 1967 and 1971, however, was highly significant (P<0.01) (Table 4). Discussion Shorebirds generally arrive earlier in the autumn at Long Point than at Kingston and with the exception of the Sanderling remain later at Kingston than at Long Point (cf. Weir and Cooke 1976). The exception of the Sanderling is probably related to its much greater abundance at Long Point as compared to Kingston. Coverage at Kingston was probably not as intense as at Long Point in July and vice versa late in the fall. Also beach pool habitat deteri- orates at Long Point as the season progresses, probably in contrast to some areas such as the sewage ponds which were censused at Kingston. Data on differential age migration at Long Point are generally more complete than the Kingston data for species which both studies cover. At Long Point, immature Semipalmated Plovers were observed about two weeks earlier in the autumn than at Kingston. Weir and Cooke (1976) found adult and immature Black-bellied Plovers at Kingston in October but at Long Point, we found only immatures. Weir and Cooke imply that Pectoral Sandpipers seen at Kingston in October were small females, but we found that at Long Point birds caught in September and October were immatures and not attributable to either sex. If breeding success or mortality in 1967 was markedly different than in 1971, either factor could account for the significant decline in shorebird abundance noted at Long Point but shorebird populations at nearby localities should have been similarly affected. There were, however, no significant differences in the num- 1977 bers of each species counted along the north- eastern shoreline of Lake Erie in 1967 and 1971 (Table 4), although the number of counts was small. Hence it seems unlikely that variable breeding success or mortality account for the differences noted at Long Point. We believe that the rising water level of Lake Erie was the most important factor causing the decline. As the lake level rose and the quality of beach pool habitat deteriorated, six of eight species that preferred to feed on beach pools declined significantly. The rising lake level did not reduce the amount of beach interface and none of the species favoring it as feeding habitat declined significantly. In fact, Sanderlings increased in numbers between 1967 and 1971. Beach pools do not occur along the north- eastern shoreline of Lake Erie and shorebirds in this area feed on mats of algae and decaying vegetation along the interface (Seeber 1966; personal observations). Rising water levels did not affect the availability of this feeding habitat and no significant changes were found in the number of shorebirds using this area. Rising lake levels should not affect the size of the breeding populations of either Killdeers or Spotted Sand- pipers at Long Point as few of either species nest on the south beach; their numbers did not decline during the study. The reasons for the significant increase in the numbers of Spotted Sandpipers from 1967 to 1971 are unclear at this time but may, in part, be due to sampling design. Piping Plovers, on the other hand, require a wide and relatively undisturbed beach for nesting (Wilcox 1959). As the water level of Lake Erie rose, a Simultaneous reduction in the width of Long Point’s south beach occurred. Since Piping Plovers nest only on the south beach, rising lake levels, by reducing the beach width between 1967 and 1971, may have been partly responsible for the decline in numbers of Piping Plovers. At least two shorebird species are somewhat traditional in their use of Long Point during the southbound migration. One percent of 731 Semipalmated Sandpipers and 7% of 471 Sanderlings banded between 1966 and 1970 were recaptured at Long Point in years subsequent to banding. The loss or deterioration of feeding habitat at a traditional staging area probably has detri- BRADSTREET ET AL: SHOREBIRDS, LAKE ERIE, 1966-71 235 mental effects on migrant shorebirds. Those birds capable of shifting to nearby staging localities may do so. Those birds incapable of shifting to a nearby locality or for which a suitable alternative does not exist would be expected to be less fit to continue migration than birds using better quality habitat. While an increase in Lake Erie water levels was observed to result in the decline, at Long Point, of numbers of shorebirds that preferred beach pools as feeding habitat, the increase resulted as part of a natural cycle in Great Lakes water levels (Laidly 1962) and the number of birds involved was very small. In certain areas of James Bay and the Maritimes where large-scale alterations of the environment are proposed, however, very large numbers of shorebirds build up fat reserves for long, non-stop flights (Hope and Shortt 1944; McNeil and Cadieux 1972; McNeil and Burton 1973). If some shorebird populations are rigid in their use of such major staging areas, and if no alternative sites exist, the destruction of those areas could destroy the populations involved. If, on the other hand, shorebirds can shift staging localities, it is important to know to what extent this may be possible, what species are behaviorally able todo so, and whether alternative areas may be capable of supporting the displaced population. More research along such lines is necessary. Acknowledgments The following volunteers of the Long Point Bird Observatory helped collect data: James Alty, Jim Bailey, Jan Bradshaw, Monica Connoly, Tim Davis, Erica Dunn, Chris Holds- worth, Geoff Holroyd, Liz Holroyd, David Hussell, Ed Keith, Anne Morris, Brian Ratcliff, Tony Salvadori, Stan Shephard, Eric Tull, and Bob Whittam. We express our warmest thanks to Long Point lightkeepers Bill Ansley, Bill Lamb, and Pete Scofield and their families; without their kindness and friendship out time at Long Point would have been much less enjoy- able. We thank A.R. Clark (Buffalo Orni- thological Society) for unpublished information and Rolph Davis, David Hussell, Guy Morrison, John Richardson, James Rising, and Lynne Stenzel for comments on the manuscript. The Long Point Company, the Canadian 236 Ministry of Transport, and the Ontario Ministry of Natural Resources granted permission to the Observatory to operate on their properties at Long Point. Financial support for general Observatory work was donated by the Canadian National Sportsmen’s show, the Federation of Ontario Naturalists, and numerous individuals. LGL Limited, environmental research asso- ciates, provided technical services in preparing this paper. Literature Cited Cottle, T. 1859. A list of birds found in Upper Canada. Canadian Record of Science 4: 231-233. Gillies, D. K. A. 1959. Winds and water levels on Lake Erie. Royal Meteorological Society, Canadian Branch 9: 12-18. Hope, C. E.and T. M. Shortt. 1944. Southward migration of adult shorebirds on the west coast of James Bay, Ontario. Auk 61: 572-576. Hussell, D. J.T. 1965. Long Point Bird Observatory — 1962 and 1963 reports. Ontario Bird Banding |: 1-44. Hussell, D. J. T.and R. D. Montgomerie. 1968. Thestatus of the Piping Plover at Long Point, 1960-1965. Ontario Field Biologist 20: 14-16. Hussell, D. J. T., D.H. Baldwin, W. A. Martin, R. D. Montgomerie, and P.S. Woodford. 1966. Long Point Bird Observatory—1964 report. Ontario Bird Banding 2: 1-51. Hussell, D. J. T., W. A. Martin, R. W. Stamp, and P.S. Woodford. 1967. Long Point Bird Observatory: 1965 report. Ontario Bird Banding 3: 30-78. Irbe, J. G. 1972. Aerial surveys of Great Lakes water temperatures—April, 1968 to March, 1970. Department of the Environment, Atmospheric Environment Service, Climatological Studies 19. 57 pp. Laidly, W.T. 1962. Regimen of the Great Lakes and fluctuations of lake levels. Jn Great Lakes Basin. Ameri- can Association for the Advancement of Science Publi- cation 71 pp. 91-105. McNeil, R. and J. Burton. 1973. Dispersal of some south- bound migrating North American shorebirds away from THE CANADIAN FIELD-NATURALIST Vol. 91 the Magdalen Islands, Gulf of St. Lawrence, and Sable Island, Nova Scotia. Caribbean Journal of Science 13: 257-278. McNeil, R. and F. Cadieux. 1972. Fat content and flight- range capability of some adult spring and fall migrant North American shorebirds in relation to migration routes on the Atlantic coast. Naturaliste Canadien 99: 589-606. Ouellet, H., R. McNeil, and J. Burton. 1973. The Western Sandpiper in Quebec and the Maritime Provinces. Canadian Field-Naturalist 87: 291-300. Page, G. and M. Bradstreet. 1968. Size and composition of a fall population of Least and Semipalmated Sandpipers at Long Point, Ontario. Ontario Bird Banding 4: 80-88. Richards, T.L. 1965. Meteorological factors affecting Great lakes water levels. Canadian Department of Transport, CIR-4182. 17 pp. Richards, T.L., J.G. Irbe, and D.G. Massey. 1969. Aerial surveys of Great Lakes water temperatures—-A pril, 1966 to March, 1968. Canada Department of Transport, Meteorological Branch, Climatological Studies 14. 55 pp. Roberts, T. S. 1955. Manual for the identification of the birds of Minnesota and neighboring states. University of Minnesota Press, Minneapolis. 738 pp. Saunders, W.E. 1926. Hudsonian Godwit in Ontario. Canadian Field-Naturalist 40: 113. Seeber, E. L. 1966. Twenty years of shorebird counts, an attempt at partial analysis. Prothonotary 32: 107-116. Seeber, E. L. 1967. The shorebird count experiment of 1967. Prothonotary 33: 117-118. Siegel, S. 1956. Nonparametric statistics for the behavioral sciences. McGraw-Hill, Toronto. 312 pp. Snyder, L.L. 1928. The Western Sandpiper, Ereuntes mauri, in Ontario. Auk 45: 207. Snyder, L. L. 1931. The birds of Long Point and vicinity. In A faunal investigation of Long Point and vicinity, Norfolk County, Ontario. III. Edited by L. L. Snyder. Transactions of the Royal Canadian Institute 18: 117-236. Townson, J. 1928. Some observations of bird life. Rod and Gun 30: 401-402. Weir, R.D. and F. Cooke. 1976. Autumn migration of shorebirds in the Kingston area of Ontario, 1964-1974. Canadian Field-Naturalist 90: 103-113. Wilcox, L. R. 1959. A twenty year banding study of the Piping Plover. Auk 76: 129-152. Received 10 February 1977 Accepted 14 May 1977 Population Dynamics, Home Ranges, and Habitat Associations of the Yellow-cheeked Vole, Microtus xanthognathus, in the Northwest Territories RICHARD J. DOUGLASS Renewable Resources Consulting Services Ltd., Edmonton, Alberta 156 0X4 Douglass, Richard J. 1977. Population dynamics, home ranges, and habitat associations of the Yellow-cheeked Vole, Microtus xanthognathus, in the Northwest Territories. Canadian Field-Naturalist 237-247. Abstract. Demographic features of Yellow-cheeked Vole, Microtus xanthognathus, populations were examined on two live- trapping grids near Chick Lake, Northwest Territories. Populations on the two grids were probably out of phase with each other. Most demographic features were similar to those described for other species of Microtus. Home range sizes were variable. Sizes were from 116 m? to 27 500 m2 and average lengths of ranges were from 15 m to 255 m. Sizes of home ranges were similar to those recorded for smaller species. Habitat data suggest broad ecological amplitude but voles occurred continuously only in dense stands of Black Spruce (Picea mariana). Micro-habitat analysis indicated association with only a few plant species and captures were associated with micro-topography. A controversy over interpretation of Pleistocene environments using fossil remains of Microtus xanthognathus as indicators is discussed. Habitat selection data suggest that some interpretations may be too restrictive. Key Words: Microtus xanthognathus, population, habitat, home range, Northwest Territories. The Yellow-cheeked Vole, Microtus xantho- gnathus, is a medium-sized microtine rodent distributed throughout the boreal forest of northwestern Canada. Little is known about its ecology, and most ecological information for this species has been contained in distribution records (Osgood and Bishop 1900; Osgood 1909; Rand 1945; Lensink 1954; Youngman 1975). These papers contain only subjective descrip- tions of populations and habitat selection. This paper presents data for Yellow-cheeked Voles obtained from an extensive live-trapping pro- gram conducted in the boreal forest in the Northwest Territories during 1973, 1974, and 1975. A secondary purpose in presenting these data stems from controversy over interpretation of Pleistocene environments using fossil remains of Yellow-cheeked Voles as indicators. Hallberg et al. (1974) questioned Guilday’s (1971) inter- pretation of cave remains as being from strictly taiga (essentially the ecotone between the boreal forest and tundra) associations, based on the presence of Yellow-cheeked Vole remains, since other mammal remains found in the same strata were considered to be more closely associated with southern boreal conditions. Data presented here on habitat selection of Yellow-cheeked Voles in the boreal forest may help clarify confusion over interpretations of Pleistocene environments. The objectives of the field work in this study were (1) to quantify demographic features of populations of Yellow-cheeked Voles and to compare these to similar parameters reported for other microtine rodents, (2) to describe sizes of home ranges of this species, and (3) to collect data concerning habitat and micro-habitat selection as inferred from live-trapping data. Methods This study was conducted in the northern boreal forest near Chick Lake, Northwest Territories. Chick Lake (65°52’ N, 128°07’ W), located west of the Mackenzie River between Norman Wells and Fort Good Hope, is approximately 70 km south of the Arctic Circle and is at an elevation of 120 m above mean sea- level. A full description of the general area surrounding Chick Lake can be found in Gubbe and Janz (1974). Sampling was conducted on two live-trapping grids and 13 live-trapping transects. Figure | shows the locations of sampling sites near Chick Lake. Data from trapping grids were used to describe demographic features of Yellow- cheeked Vole populations, home range sizes, and micro-habitat selection. Data from transects 23), 238 THE CANADIAN FIELD-NATURALIST CHICK FIGURE 1. Map of the study area showing the locations of trapping grids and transects. Numbers indicate the locations of the following sampling sites: 3, Creek bank; 4, small lakeshore; 5, burn; 6, birch forest; 7, open spruce; 8, shore of Chick Lake; 9, willow bar; 10, hillside; 11, Alder-covered lakeshore; 12, forested lakeshore; 13, White Spruce forest; 14, river bank; 15, hilltop; 16, Black Spruce forest. were used to describe distribution of Yellow- cheeked Voles in relation to general habitat types occurring in the study area. Data collected by McDonald (1974) in “a burned over” area near Chick Lake are also included in the habitat discussion. The two live-trapping grids were constructed on opposite sides of Chick Lake. Each grid consisted of 250 stations with one trap at each, marked with surveyor’s stakes spaced at 15-m intervals. The grids were 10 columns wide by 25 rows long and encompassed areas of 4.86 ha each. Sampling was conducted during July through October 1973, June through October 1974, and during June, August, and September 1975. The grids were trapped for the first 10 days of each month during 1973 and 1974 except October, which was sampled for 7 days, and during the first 7 days of each month during 1975. One hundred traps were placed on each transect at 8-m intervals and were arranged in either a grid or line. Traps were set on each transect for 3 days each month. Sherman live-traps, 23 X 8 X 9 cm, were used on both grids and transects during all but the first month of the study. During the first month, home-made can traps were used. Traps were baited with rolled oats, covered with moss, and checked twice daily. Each animal was individually numbered by toe-clipping and was released at the point of capture. The number, location of capture, sex, breeding condition, and weight were recorded for each animal every time it was captured. Quadrats (one per trap station) of 1 m* were used to determine frequency of occurrence and percentage cover for all plant species on both grids. Coverage values were classified according to a scheme described by Daubenmire (1959). The degree of micro-relief or micro-topography within a 2-m radius of each trapping station was classified according to the following designa- tions: class 0, ground flat; class 1, small NOT. hummocks up to 1I5cm tall; class 2, tall hummocks >15 <30 cm tall; class 3, undercut and collapsing hummocks >30 cm and <50 cm tall; class 4, areas with hummocks >S50 cm tall. Results and Discussion Population The Yellow-cheeked Vole comprised only a small portion of the small mammal community as indicated by live-trapping in the Chick Lake area. In 102300 trap-nights (TN) of effort, nine species of small mammals were captured. These were Northern Red-backed Voles (Clethri- onomys rutilus), 10319 captures of 2345 individuals; Yellow-cheeked Voles (Microtus xanthognathus) 937 captures of 360 individuals; Meadow Voles (M. pennsylvanicus), 863 cap- tures of 479 individuals; Masked Shrew (Sorex cinereus), 30 captures of 30 individuals; Nor- thern Bog Lemming (Synaptomys borealis), 25 captures of 23 individuals; Heather Vole (Phenacomys intermedius), 15 captures of 8 individuals; Red Squirrel (Tamiasciurus hud- sonicus), 12 captures of 10 individuals; Ermine (Mustela erminea), 3 captures of 3 individuals; and Meadow Jumping Mouse (Zapus hud- sonius), | capture of | individual. No previous studies have described the demographic features of Yellow-cheeked Vole populations. Any mention of population size in taxonomic reports have not been quantitative, although apparently Yellow-cheeked Voles can be fairly numerous and fluctuate in number. Rand (1945) suggested that these voles were subject to violent population fluctuations. Lensink (1954) interviewed fur trappers who indicated that the voles were numerous, and natives at Old Crow in the Yukon Territory said that these voles exploded in numbers about every 20 years (Youngman 1975). In this study, sizes of populations of Yellow- cheeked Voles are represented by three mea- sures: minimum number of individuals known to be present per grid per month, the number of individuals per hectare per month (number present/area of grid), and the number of captures per 100 TN per month (Figure 2). Minimum densities were less than I/ha and maximums were near 10/ha. As recommended by Krebs (1966) no attempt was made to estimate population density by capture-recap- DOUGLASS: MICROTUS XANTHOGNATHUS IN NWT 2B ture techniques because of non-random sam- pling. A sharp decrease in the population occurred on Grid B between July and August 1973 and was almost entirely the result of trap mortality. Yellow-cheeked Voles were too large for the home-made can traps and many were killed by the closing doors of traps. No animals were directly killed by traps after the can traps were replaced by Sherman traps. A sharp decrease in numbers on both grids, which occurred between September and October 1974, may have been the result of inhibited activity in October caused by snow depths of 30cm and temperatures of -33°C. Even though weather conditions were not as severe during October 1973, population indices for October of both years are not considered as accurate as they may have been during other months. Yellow-cheeked Vole populations on the two grids appeared to be out of phase with each other in their yearly fluctuations. The population on Grid A went through two summers (1973, 1974) of relatively low numbers, increasing slightly over winter between these summers and then increased over the 1974-1975 winter, thus entering an apparent increase phase with larger numbers occurring during 1975. The popula- tion on Grid B was in a high or increase phase when the study began in 1973, was accidentally reduced, increased over winter, and reached a peak phase in 1974 followed by a decrease that lasted through the winter with the population remaining very low through 1975. Essentially populations on the two grids were fluctuating in opposite fashion. Two patterns of population fluctuation were observed, seasonal and multi-annual. Within each of the multi-annual population fluctua- tions there was a seasonal change tht occurred on both grids. Populations were generally small early in the year, increased through the summer, and began to decline as fall approached. Thus while the seasonal fluctuations were similar on the two grids, the multi-annual fluctuations were asynchronous. Krebs and Myers (1974) describe the “demographic machinery” which can cause fluctuations as being comprised of reproduction, mortality, and dispersal. Some data concerning each of these were collected for Yellow-cheeked Voles at Chick Lake. 240 THE CANADIAN FIELD-NATURALIST Vol. 91 @Grid A 4Grid B 10 9 S$ 40 8 q 7 © 30 6 z 5 = S fe) 5 20 42 A= 3 = = 10 2 1 > ips) Captures/100 Trap Nights = ww O J J J A S) *traps open for 7 days only 1975 FIGURE 2. Population sizes of Yellow-cheeked Voles on two trapping grids near Chick Lake, Northwest Territories, represented by the number of individuals captured and the number of captures per 100 trap nights. As first described for M. agrestis by Chitty (1952) large individuals occur more often in high populations than in low populations. A similar situation is evident for Yellow-cheeked Voles as indicated by the distribution of live weights of males (Figure 3). Voles weighing 90 g or more were trapped only during high population densities on each grid, which is consistent with the general microtine population syndrome. Reproduction Data concerning pregnancy rates, percentage of animals in breeding condition, length of breeding season, and sex ratios were collected for Yellow-cheeked Voles. Figure 4 shows the percentage of female Yellow-cheeked Voles that were apparently pregnant, and the percentage of adults in breeding condition (vulva swollen or vaginal membrane absent for females; testes in scrotal position for males). The sample was not sufficient to permit a meaningful assessment of trends, but there was no indication of increased pregnancy rates preceeding or accompanying the population increases or peaks demonstrated by Yellow-cheeked Voles. The sample size was again too small to show any association between percent in breeding condition and population sizes or increases. Although sample sizes are small, the indication from the data that neither pregnancy nor breeding rates increased with or prior to population increases is consistent with changes in similar parameters described for Live weight in gms. Live weight in gms. J A S O DOUGLASS: MICROTUS XANTHOGNATHUS IN NWT 241 1974 June A S FIGURE 3. Live weights of male Yellow-cheeked Voles on two trapping grids near Chick Lake, Northwest Territories. Each square represents the weight of one vole. other microtine rodents as discussed by Krebs and Myers (1974). Since sampling did not begin until June during each season the onset of breeding can only be inferred from the date the first apparently juvenile voles (less than 30 g) appeared. Voles this size were not captured before July, suggesting that breeding occurred in May (Figure 3). The end of the breeding season, however, occurred by October on both grids during 1973 and 1974 (Figure 4). If this is an accurate assessment, there is no indication that the breeding season was longer during the possible increase or peak phases that occurred on Grid B during 1973 and 1974 or on Grid A during 1975. Other microtines generally have increased lengths of breeding seasons prior to peak phases (Krebs and Myers 1974). Changes in sex ratios of microtines have not been found to be associated in any consistent way with population fluctuation (Krebs and Myers 1974). Sex ratios were calculated for Yellow-cheeked Voles on both grids for each trapping season. The male to female ratios were as follows: 1973 1974 1975 Grid A 20:28 24:24 34:29 Grid B 29:58 34:72 27 The ratios appear to have been associated more with a particular grid than with population density. Ratios on Grid A were close to 50:50 and those on Grid B favored females. As described for other microtines, changes in sex ratio did not appear to be associated with population fluctuations. Mortality and Dispersal The inverse of mortality, survival, will be considered since there was no way of knowing 242 7, Pregnant 1973 7, in Breeding Condition THE CANADIAN FIELD-NATURALIST Vol. 91 Grid A “Grid B 1974 1975 A Ss O J J A ) FIGURE 4. Breeding status of Yellow-cheeked Vole populations on two trapping grids near Chick Lake, Northwest Territories. whether voles died in situ or emigrated when they ceased being captured on the grids. Figure 5 shows the survival rates of voles on both grids. Survival on Grid A was moderate to low during low population phases 1973 and 1974 but was high during the increase phase, 1975. Grid B population survival rates were somewhat higher than those on Grid A during 1973 and 1974, during high or increased population phases on Grid B, but went to zero during the low phase in 1975. Survival was high in populations that were in an increase or peak phase and was low during phases of low population densities. Krebs and Myers (1974) described similar phenomena for other microtines. Only anectodal information is available concerning dispersal of Yellow-cheeked Voles. In this study, voles were captured in more different habitats, i.e., Black Spruce (Picea mariana) forest, small lakeshore, alder (A/nus crispa)-covered lakeshore and the river bank on the south side of Chick Lake during 1973 (Table 1) when populations were higher on that side of the lake, as indicated by populations on Grid B, than during 1974 or 1975. Also, Yellow-cheeked Voles were captured in an additional habitat, open spruce, as well as the dense Black Spruce forest on the north side of the lake during 1975 when populations were increasing on that side as indicated by populations on Grid A. If the occurrence in several habitats can be considered to be indicative of dispersal, the dispersal during high population densities, and particularly increasing densities, coincides with dispersal phenomena described in the population syndrome of Krebs and Myers (1974). IVY 90 1973 1974 80 70 60 50 40 30 7, Montly Survival 20 10 J A S O J J A DOUGLASS: MICROTUS XANTHOGNATHUS IN NWT 243 ® Grid A 4Grid B 1975 iS) O J A S FiGuRE 5. Monthly survival rates for Yellow-cheeked Voles on two trapping grids near Chick Lake, Northwest Territories. Survival is expressed as the percentage of voles recaptured during month T + | that were originally captured during month T. Home Range Sizes of home ranges were calculated using the convex polygon index (Jenerich and Turner 1969) for each Yellow-cheeked Vole captured three or more times. The maximum lengths of home ranges were also calculated by measuring the distance between the two most distant capture points for each individual captured three or more times. Home ranges were not calculable when the only three capture points formed a line. Maximum lengths of home range were cal- culated for these, however. During the three summers a total of 29 home range sizes and 42 lengths of home ranges were calculated for Yellow-cheeked Voles. Table 2 shows the average sizes and lengths of home ranges according to grid, year, and sex. Sample sizes were insufficient to make any conclusions from number of captures and home range size for this species. Average measure- ments of both parameters were extremely variable. Average home ranges varied from 116 m? for males on Grid A during 1973 to 2671 m2? for females on Grid B during 1974. The smallest average maximum length of home range was 25 m for males on Grid A during 1973 and the largest was 18! m for females on Grid B during 1973. The only other home range sizes previously reported for Yellow-cheeked Voles were for two individuals found near San Sault rapids in the Northwest Territories (Mitchell 1973). The home ranges were 769 m2 for a male and 405 m? for a female, both of which fall within the ranges of those measured at Chick Lake. Yellow- cheeked Voles also had home ranges similar in size to those previously recorded for the smaller Meadow Vole. Some of the recorded home ranges of Meadow Voles are 232 m2 (Hamilton 1937); 809 m2? to 2023 m? (Blair 1950); and 267 m2 to 2023 m2 (Hayne 1950); and 232 m? to 8829 m2 at Chick Lake (Douglass 1975). Ap- parently the relatively large Yellow-cheeked Vole does not require home ranges larger than those normally required by smaller voles. Habitat Associations Habitat selection was inferred from associa- 244 THE CANADIAN FIELD-NATURALIST Vol. 91 TABLE 1—Description of live-trapping sites near Chick Lake, Northwest Territories, TN = trap nights Title Topography Dominant vegetation Captures/100 TN 1973 1974 1975 Sites associated with “edges” Creek bank Thermally and hydro- Black Spruce, lichens, 0.0 0.0 0.0 logically eroded, very moss steep, broken Small lakeshore Gently sloping, flat Sedges, willows 0.1 0.0 0.0 Shore of Chick Lake Gently sloping, flat Sedges, grasses 0.0 0.0 0.0 Alder-covered lakeshore Gently sloping Alders, grasses 0.1! 0.0 0.0 Forested lakeshore Steeply sloping Black Spruce, White Spruce, 0.0 0.0 0.0 . mosses River bank Cutbanks of river, Alders, Black Spruce, sedges 0.1! 0.0 0.0 boulder covered, steep and slumping Sites not associated with “edges” Hilltop, Grid A and B Level and hummocky Black Spruce, Ledum, sedges, 1.6 1.6 1.13 lichens Black Spruce forest Gently sloping with Black Spruce, lichens, mosses 0.0 0.0 0.0 small hummocks Burn* Gently sloping with Burned in 1969, standing dead 0.2 0.0 = large hummocks trees, bog birch, willows and sedges Birch forest Level to gently sloping Birch trees, Ledum 0.0 0.0 0.0 Open spruce Level and flat Sparse Black Spruce, lichens 0.0 0.0 0.12 Willow bar Level and flat, river Alders, willows 0.0 0.0 0.0 bank Hillside Steep, slumping Birch trees, alders 0.0 0.0 0.0 hillside White Spruce forest Level, flat Mature White Spruce, little 0.0 0.0 0.0 understory *Sampled by McDonald (1975) during 1973 and 1974. 'Represents single captures made during August 1973. Represents single capture made during September 1975. 3Captures occurred at these locations during every sampling period. tion of vole captures with general habitat type and micro-habitat as represented by individual plant species and micro-topography. It was assumed that habitats in which captures of Yellow-cheeked Voles occurred were more suit- able for the voles than those habitats without captures. Table | lists the sites that were sampled, gives a general description of each, and average captures per 100 TN for each. More detailed information can be found in DL suglass (1975). Of the 16 sites sampled, Yellow-cheeked Voles were captured in only eight: Grids A and B plus the hilltop, burn, open spruce, small lakeshore, alder-covered lakeshore, and river bank tran- sects. Yellow-cheeked Voles were captured all three years only in non-edge habitats dominated by Black Spruce (both grids and the hilltop transect). The failure to capture Yellow-cheeked Voles in a fourth Black Spruce habitat (Black Spruce forest, Table 1) indicates that the Black Spruce forest was sufficient for continuous occupation but was not necessarily always occupied. Captures were not made during every sampl- ing period in most habitats (Table 1). It is hypothesized that these habitats were marginal, being occupied by transients, as suggested for Muskrats by Errington (1967) when “optimal” habitats, presumably Black Spruce forests, had peak populations or as the populations in the “optimal” habitats were increasing (Krebs and Myers 1974). Guilday et al. (1964) reviewed the literature on 1977 DOUGLASS: MICROTUS XANTHOGNATHUS IN NWT 245 TABLE 2—Average sizes (mean + 1 SD) of home ranges (m°) and maximum lengths (m) of range for Yellow-cheeked Voles near Chick Lake, Northwest Territories, during 1973, 1974, and 1975 1973 1974 1974 Grid A (N) GridB (N) Grid A (N) Grid B (N) Grid A (N) Grid B (N) Size of Home Range Males 116 — (1) 522+487 (2) 754+1209 (6) 1161 +957 (4) 1433 + 1375 (3) = (0) Females = @ Wol—— @) LY] — @) Alea ©) 24 se 715 G) = (0) Maximum Length of Home Range Males 25-3 (2) 37] = 30) 24(5) 1S 3is=4 1 (4) 44+50 (4) 82+37 (3) = (0) Females 29 rea ia (2) eal 8 lis=t=19 7], (2) et Sy =4 Oem) en 59) = 49) (O73) td (6) = (0) Yellow-cheeked Vole habitats and concluded that this species was adapted to a broad range of habitats. Youngman (1975) examined both pub- lished and unpublished accounts of Yellow- cheeked Vole habitats and also concluded that this species displayed broad ecological ampli- tude but found that most records indicated association with recently disturbed areas near mineral soils. The habitat data from Chick Lake also suggest a broad ecological amplitude in that Yellow-cheeked Voles were captured in several different habitat types. The Chick Lake data indicate, however, that even though Yellow- cheeked Voles could be captured at one time or another in many different habitats, including a recently burned area and river bank with mineral soils, they could be captured consistently only in the Black Spruce forest which was relatively undisturbed and situated on organic soils. The first step in describing micro-habitat selection was the determination of whether frequency of occurrence of plant species at trapping grid stations where voles were captured was different from the overall frequency of occurrence of all plant species on each grid. A goodness of fit chi-square analysis (Sokal and Rolf 1969) was used to compare expected fre- quencies at capture points to observed fre- quencies. For each capture, the vegetation plot recorded for the station where the capture occurred was included as an individual observa- tion in the analysis. All micro-habitat analyses were performed by computer on plant and capture data collected during 1974. A summary of the vegetational characteristics of both grids is shown in Table 3. The chi-square analysis showed that Yellow- cheeked Vole captures occurred at stations with frequencies of occurrence of all plant species that were significantly different from the frequency of occurrence of all plant species as they occurred on the grid, on Grid A Sx*° = 73.37 df =37 P<0.05 but not Grid B Yx* = 42.24 df= 38 P>0.05. This suggests that Yellow- cheeked Voles were being selective of areas in which they were captured on Grid A but not Grid B. The analysis was carried further to determine which individual plant species or locations in which they occurred were selected for or against. Chi-square association tests (Cole 1949) were employed to determine whether there was a TABLE 3—Summary of soil and vegetational characteristics of Grid A and Grid B at Chick Lake, Northwest Territories Grid A Grid B Soil characteristics (mean values) (depth in cm) Organic layer 37 19 Active layer 55 48 Tree characteristics (mean values) Height (m) 4.3 4.3 DBH (cm) 5.0 3.8 Density (stems/ ha) 6869.0 8714.0 Plant cover* and number of species % Cover No. species % Cover No. species Trees 13.4 6 23.1 4 Shrubs 79.2 10 67.0 7 Herbs DRS 7 22D, 10 Mosses 68.3 6 56.8 6 Lichens 32.4 8 39.1 11 Litter {| — 4.8 — *See Daubenmire (1959). 246 relationship between captures of Yellow- cheeked Voles and the occurrence of any par- ticular plant species at capture points. Cor- relation coefficients were also calculated be- tween the number of captures/ 100 TN and the percent coverage of each plant species. Table 4 presents the results of these analyses only for those species found to be statistically signifi- cantly associated with or correlated with vole captures. Fourteen species and one major group, Gramineae, were found to be associated (sig- nificant chi-square association values) with or correlated (significant correlation coefficients) with Yellow-cheeked Vole captures on either or both of the two grids. Captures were determined to be significantly associated with 11 species plus Gramineae, and the number of captures/ 100 TN showed correlations with cover of three species and Gramineae. Captures on both grids were associated with the presence of Vaccinium uliginosum (nega- THE CANADIAN FIELD-NATURALIST Vol. 91 tively), Blueberry; Picea mariana, Black Spruce; and captures on both were associated with and correlated with Gramineae. These two species and Gramineae should probably be considered to have been the most important to Yellow- cheeked Voles. Gramineae, Ca/amogrostis canadensis in par- ticular, was considered as food for Yellow- cheeked Voles by Youngman (1975), and cap- tures were associated with the presence and amount of Gramineae in this study, possibly because the voles were attracted in search of food. Small P. mariana less than 2 m tall usually formed very dense cover on the two grids. The association between captures and the occurrence of this species was probably indicative of the use of small P. mariana as cover. The negative association between captures and the occurrence of V. uliginosum is puzzling since Youngman (1975) found berries from this species in the mouths of snap-trapped Yellow-cheeked Voles, TABLE 4— Plant species associated with Yellow-cheeked Vole captures on two live-trapping grids near Chick Lake, Northwest Territories during 1974. Chi-square values were calculated from frequency of occurrence of plant species and frequency of capture of voles. Correlation coefficients were calculated from cover values of plant species and numbers of captures/ TN of Yellow-cheeked Voles Grid A Grid B Chi Chi- square r df square r df Vascular plants > 0.5 m < 2 m in height Alnus crispa 12.083* —).389 18 0.135 +0.056 38 Betula glandulosa 5.099* +0.663 7 0.002 —0.155 17 Ledum groenlandicum 4.024* —).078 32 0.236 —0.051 92 Potentilla fruticosa 7.910* +0.306 20 0.123 —).022 40 Rosa acicularis 3.74 +0.205 14 4.387* —0.134 44 Vaccinium uliginosum 13.240* —0.001 34 4.051* —0.097 84 Picea mariana 9.071* +0.192 41 4.931* +0.151 76 Vascular plants < 0.05 m in height Empetrum nigrum 0.028 +0.745* 6 0.915 — = Salix sp. 1.199 —0.152 14 3.805 +0.919* 3 Gramineae 3.978* +0.391* 28 10.987* +0.033 44 Linnaea borealis 4.150* +0.971 l 0.410 —).119 27 Lichens Cladonia mitis 6.088* +0.001 5 0.013 —0.130 66 Peltigera sp. 18.277* +0.400 16 3.524 —0.144 60 Liverworts Hedwigia sp. 0.278 Talal 19 1.185 +0.236* 78 Unknown species — a 9.086* +0.127 SY +< 2 m in height. *P<0.05. 1977 and several voles were observed eating these berries near Chick Lake. It may be that demon- strations of habitat selection via trapping can be inaccurate and possibly in this situation com- pletely opposite from the actual phenomenon. Results of the micro-topography analyses suggested that micro-topography was probably more important to Yellow-cheeked Voles than were individual plant species. Correlation coef- ficients calculated for captures/100 TN versus degree of micro-relief for Grids A and B were +0.967 P< 0.05 and +0.981 P< 0.05, respec- tively. Greater micro-relief probably increased the amount of escape cover for these voles, especially in a forest with a moss- and lichen- dominated understory which provided little herbaceous escape cover. Acknowledgments Data for this paper were gathered while field studies were being conducted for Canadian Arctic Gas Study Limited by Renewable Re- sources Consulting Services Ltd. I thank R. M. Moore and C. J. Krebs for critically reviewing the manuscript; K. S. Douglass, L. Fisher, B. Wooley, A. McNaughton, M. Mair, and G. Klassen for aiding in the field work; D. Gubbe and D. Burr for assisting in the analysis of vegetation; and G. Lance for providing com- puter expertise. Literature Cited Blair, W. F. 1940. Home ranges and populations of the Meadow Vole in southern Michigan. Journal of Wildlife Management 4: 149-161. Chitty, D. 1952. Mortality among voles (Microtus agrestis) at Lake Vyrnwy, Montgomeryshire in 1936-9. Philo- sophical Transactions Royal Society of London, Series B. 236: 505-552. Cole, L. C. 1949. Measurement of interspecific association. Ecology 30: 411. Daubenmire, R. F. 1959. A canopy coverage method of vegetation analysis. Northwest Science 33: 43-64. Douglass, R. J. 1975. A study of the ecology of small mammals. /n Second preliminary report of monitoring conducted at Chick Lake, N.W.T. Edited by D. Reid. Unpublished, prepared by Northern Engineering Services Company Limited for Canadian Arctic Gas Study Limited, Calgary. 70 pp. Errington, R. L. 1967. Of predation and life. Iowa State University Press, Ames. 277 pp. Gubbe, D. and A. Janz. 1974. Landscape classification of the Chick Lake area. /n Preliminary report of monitoring conducted at Chick Lake, N.W.T. Edited by D. Reid. DOUGLASS: MICROTUS XANTHOGNATHUS IN NWT 247 Unpublished, prepared by Northern Engineering Services Company Limited for Canadian Arctic Gas Study Limited, Calgary. 103 pp. Guilday, J. E. 1971. The Welsh Cave peccaries (Platy- gonus) and associated fauna. Kentucky Pleistocene Annals of the Carnegie Museum 43: 249-320. Guilday, J. E., E. W. Hamilton and A. D. McCrady. 1964. New Paris Number 4: A Pleistocene cave deposit in Bedford County, Pennsylvania. National Speleology Society Bulletin 26: 121-194. Hallberg, G.R., H. A. Semken and L.C. Davis. 1974. Quaternary records of Microtus xanthognathus (Leach) the Yellow-cheeked Vole, from Northwestern Arkansas and Southwestern Iowa. Journal of Mammalogy 55: 640-645. Hamilton, W. J. 1937. Activity and home range of the Field Mouse (Microtus pennsylvanicus). Ecology 18: 255-263. Hayne, D. W. 1950. Apparent home range of Microtus in relation to distance between traps. Journal of Mam- malogy 31: 26-39. Jenerich, R.I. and F.B. Turner. 1969. Measurement of non-circular home range. Journal of Theoretical Biology 22: 227-237. Krebs, C. J. 1966. Demographic changes in the fluctuating populations of Microtus californicus. Ecological Mono- graphs 36: 239-273. Krebs, C.J. and J. H. Myers. 1974. Population cycle in small mammals. Advances in Ecological Research 8: 267-399. Lensink, C. J. 1954. Occurrence of Microtus xanthognathus in Alaska. Journal of Mammalogy 35(2). McDonald, D. 1975. Effects of fire on small mammals in the Northwest Territories. M.Sc. thesis, Brock University, St. Catharines. 73 pp. Mitchell, S. 1971. An investigation to postulate the re- action of Northwest Territories small mammals to vege- tation covering the proposed Mackenzie Valley Gas Pipeline. Edited by D.L. Dabbs, W. Friesen, and S. Mitchell. Canadian Arctic Gas Study Limited Biological Report Series, Volume 2. 65 pp. Mohr, C.O. 1947. Table of equivalent populations of North American small mammals. American Midland Naturalist 38: 223-231. Osgood, W. H. 1909. Biological investigations in Alaska and Yukon Territory. North American Fauna 30. Osgood, W.H. and L.B. Bishop. 1900. Results in bio- logical reconnaissance of the Yukon River Region. North American Fauna 19. Rand, A. L. 1944. Mammal investigations on the Canol Road, Yukonand Northwest Territory. National Museum of Canada Bulletin. 99 pp. Sokal, R. R. and F. J. Rohlf. 1969. Biometry; principles and practice of Statistics in biological research. W. H. Freeman and Company, San Francisco. 776 pp. Youngman, P. M. 1975. Mammals of the Yukon Territory. National Museum of Canada Publications in Zoology, Number 10. pp. 98-101. Received | February 1977 Accepted 18 June 1977 Nesting and Brood Ecology of Lesser Scaup at Waterhen Marsh, Saskatchewan JAMES E. HINES Department of Biology, University of Regina, Regina, Saskatchewan S4S 0A2 Hines, James E. 1977. Nesting and brood ecology of Lesser Scaup at Waterhen Marsh, Saskatchewan. Canadian Field- Naturalist 91(3): 248-255. Abstract. The nesting and brood ecology of the Lesser Scaup (Aythya affinis) was observed at Waterhen Marsh in central Saskatchewan. Most scaup nests were initiated during the first two weeks of June and the peak of hatching occurred during the middle two weeks of July. The mean clutch size was 9.70 + SE 0.21 (n = 56); if clutches of 13 and 14 are omitted, the mean becomes 9.47 + 0.18 (m = 53). Several incidents of egg parasitism involving scaup were noted. Insular, well-concealed nest sites featuring plants (especially grasses) in the 21- to 60-cm height range were often selected. Twenty-eight (76%) of 37 nests were successful. Striped skunks (Mephitis mephitis) and Common Crows (Corvus brachyrhynchos) caused most of the nesting failures. The high social tolerances exhibited by brooding scaup and the concentration of broods in areas of suitable habitat resulted in the formation of mixed broods. Redhead (Aythya americana) ducklings frequently joined these broods. Possible advantages of this créching are discussed. Although the Lesser Scaup (A yvthya affinis) is one of the most abundant ducks in North America, much remains to be learned about its spring and summer biology. Earlier studies dwelt on the life history of the species (Bent 1923; Kortright 1942; Gehrman 1951). More recently, food habits have been investigated (Rogers and Korschgen 1966; Dirschl 1969; Bartonek and Hickey 1969; Bartonek and Murdy 1970; Sugden 1973; others), as have features of the scaup’s breeding biology (Rogers 1959, 1964; Dwernychuk 1968; Long 1970; Trauger 1971). Previous authors have referred to the Lesser Scaup’s habit of forming mixed broods that are often led by several hens (Munro 1941; Hoch- baum 1944). A similar type of behavior has been frequently observed in scoters (Melanitta fusca), eiders (Somateria mollissima), and Shelducks (Tadorna tadorna) and has been interpreted asa form of créching or communal “baby-sitting” of the young (Koskimies 1955; Hildén 1964; Hori 1964; Ahlen and Andersson 1970; Gorman and Milne 1972). This differs from the formation of “giant broods” led by a single hen, typical of some species of the tribe Mergini (Hildén 1964). During a 2-year study of the ecology of the Gadwall (Anas strepera), | had the opportunity to observe some aspects of the nesting and brood ecology of the Lesser Scaup. As mixed broods of scaup and Redheads (Aythya americana) were frequent, this phenomenon was also investi- gated. Study Area Waterhen Marsh (50°51’ N, 105°02’ W) is located 8 km south of the town of Kinistino in the aspen parkland of Saskatchewan. The marsh was drained in the 1920s, but after agricultural attempts failed and a fire broke out in the underlying peat soils the basin was reflooded. An earthen dam, constructed in 1938 by Ducks Unlimited (Canada), aids in control of water levels in the 1530-ha impoundment (Figure 1). The marsh basin is very shallow, exceeding a depth of | m only in the drainage ditches. The underlying soils have remained quite un- consolidated since the fire and this has pre- vented the establishment of rooted submergent vegetation in most portions of the marsh. Only in the northeast corner of the marsh are submergents, primarily spiked water milfoil (Myriophyllum exalbescens), abundant. Emer- gent vegetation, especially cattail (Typha Jati- folia), is a dominant feature of the marsh, covering 30-35% of the total area. Large bays of open water are separated by the cattail stands. Dense growths of cattails, sedges (Carex spp.), bulrushes (Scirpus spp.), and whitetop grass (Scolochloa festucacea) are present in the northwestern and southeastern corners of the basin. 248 1977 HINES: LESSER SCAUP ECOLOGY, SASKATCHEWAN 249 emergents = J willows @ island b brood area oO 100 200 meters eae Oo 2000 feet FiGuRE |. Waterhen Marsh, drawn from a 1970 aerial photo. Throughout the marsh there are a great many the invertebrate fauna are the benthic midge or aquatic invertebrates, which are an important chironomid larvae and free-swimming clado- source of food for waterfowl; two members of cerans. 250 There were three types of habitat available for ground-nesting ducks at Waterhen Marsh: upland areas, ditchbanks, and islands. Ten rectangular artificial islands, each approxi- mately 10X 30m in dimension, were con- structed prior to the reflooding of the marsh and these, along with a 2.2-ha natural island, constitute most of the island habitat. Many of the ditchbanks have been substantially eroded by waves and are highly segmented and insular in nature. Methods Field work was conducted from the second week of May to the first week of September in 1972 and 1973. Nests were located by systematic inspections of the islands and part of the ditchbanks, and by less intensive searches of the upland areas. Most nests were marked, usually at a distance of 10 m away, with a small strip of red surveyors tape. Other nests were relocated by topographic and vegetational features only. I tried to recheck nests once or twice before hatching to gather information on date of nest initiation and clutch size. Whenever possible, date of nest initiation, date of hatching, clutch size, distance from water, and fate of the nest were recorded. The plant species, canopy coverage, and height of vegetation at the nest site were also recorded. Canopy coverage above the nest was estimated to belong to one of six classes, following Daubenmire (1959): 0-5% canopy coverage, 5-25%, 25-50%, 50-75%, 75-95%, and 95-100%. Brood observations were made from a canoe or motorboat, or by scanning the marsh from the uplands, ditchbanks, or islands. Use of 7- or 8- power binoculars and a 40-power spotting scope facilitated these procedures. The age of duck- lings (see Gollop and Marshall 1954), the number of ducklings, the features of the habitat, and behavior of the broods were noted. When mixed broods were encountered, I tried to identify the ducklings to species, age, and number. Results and Discussion Nesting Chronology and Clutch Size Lesser Scaup were among the most numerous ducks on the study area and were already present at the commencement of field studies m both THE CANADIAN FIELD-NATURALIST Vol. 91 years. Many or all of these early migrants may have moved further north to nest (Trauger 1971). The peak of nest initiation did not occur until the first two weeks of June. The most frequently occurring clutches were 10, 9, and 8 eggs respectively (Figure 2). If clutches of more than 14 eggs are considered as parasitized (Hildén 1964), the average clutch size is 9.70 + SE 0.21 eggs (n = 56). If 12 eggs are considered as the maximum clutch (Weller 1959; Weller et al. 1969), the average is 9.47+0.18 eggs (n = 53). Compound clutches or dump nests, pre- sumably resulting from two or more scaup hens laying eggs in a single nest, contained 16, 16, 17, 18, and 19 eggs, respectively. The habit of parasitic egg-laying has been frequently noted for this species and is known to occur both intra- and inter-specifically (Weller 1959; Vermeer 1968; Long 1970). At Waterhen Marsh, scaup eggs were frequently found in Gadwall nests, 26 of 295 completed Gadwall first clutches being parasitized. One scaup egg was found in each of 18 (69%) of the 26 Gadwall nests but as many as five eggs were present in one nest. The average first clutch size for Gadwall nests parasitized by scaup was 11.0 + 0.3 Gadwall eggs as compared with 10.4+0.! for all normal first clutches. Therefore, egg parasitism did not reduce the average clutch size of the host duck. Weller (1959) and Joyner (1976) have shown that the clutch of the host species can be reduced in such instances. Three scaup clutches of 9, 10, and 9 eggs found on the large natural island in 1973, contained 8, 6, and 6 Gadwall eggs respectively. As few other scaup clutches were found to contain Gadwall eggs and, as other studies show little evidence of a high rate of egg parasitism by Gadwalls, it is possible that these nests were initiated by Gadwalls and later taken over by scaup. Noeggs of other duck species were found in scaup nests. Nest-site Selection and Nesting Success Although the nests of many dabbling ducks were found up to 500 m from the marsh edge, all scaup nests were found close to water. More than 50% of the nests were situated within 5 m of the water’s edge and approximately 75% were within 10 m. Nest sites were usually dry and at least 30 cm above water. Many of the ditchbank 2 aa r NN O 13 E 20 12 \ oy O Li. O _ es NXX, HINES: LESSER SCAUP ECOLOGY, SASKATCHEWAN 251 NN 7 8 9 10 11 12 13 14 15 16 17 18 19 CLUTCH SIZE FiGURE 2. A frequency distribution of Lesser Scaup clutch sizes. The numbers of clutches are indicated above the histogram. nests were situated right along the edge of the bank, within |m of water, in thick gram- inaceous (grassy) vegetation. In contrast to dabbling ducks’ nests, only 3 of 64 scaup nests were situated in upland areas, 18 were on the ditchbanks, and 43 were on the islands. As the ditchbanks were much divided by erosion, most of the nests there were essentially insular in nature. No nests were found among emergent plants, but these areas were not intensively investigated. Scaup often used graminaceous cover for nesting (Table 1). Awnless brome (Bromus inermis) was the most common cover species, providing the greatest percentage of the canopy at 35% of the nest sites. Western snowberry (Symphoricarpos occidentalis) was the domi- nant at 19% of the nests, all on the large natural island. The only forb which was dominant at more than one nest was Canada thistle (Cirsium arvense). Overall graminaceous plants were TABLE 1—The dominant plant species at 54 Lesser Scaup nests at Waterhen Marsh Nests Species Growth form Number (%) Bromus inermis graminaceous 19 ( 35) Symphoricarpos occidentalis shrub 10( 19) Carex spp. graminaceous Fe Ms) Scolochloa festucacea graminaceous 6( 11) Cirsium arvense forb 6( 11) Miscellaneous grasses graminaceous 40 7) Axyris amaranthoides forb iG) Rosa woodsii shrub C2) Total 54 (100) U2 dominant at 67% of 55 nests, shrubs at 20%, and forbs at 13%. The limited use of forbs as nesting cover 1s somewhat surprising, as plants of this physio- gnomy were an important cover species else- where (Dwernychuk 1968; Long 1970). At Waterhen Marsh, the stands of forbs such as common nettle (Urtica gracilis) and Russian pigweed (A xyris amaranthoides) appeared to be too tall to be readily used by scaup. Nesting females had great difficulty in getting airborne from these patches and apparently for this reason used mainly the edges of such cover. Most scaup nests were situated in cover in the 21- to 60-cm height range (Table 2). Vegetation of 20cm or less in height was avoided, presumably because it provided poor conceal- ment. Nests in the taller height classes (> 60 cm) were normally located near the edge of the stands of vegetation as discussed above. Long (1970) found that Lesser Scaup preferred to nest in vegetation 15 to 34 cm in height. He believed that this resulted from the selection for cover which provided adequate concealment, yet did not too greatly obscure the hen’s view of the surrounding area. At Waterhen Marsh, Lesser Scaup used decidedly taller vegetation than indicated in Long’s study. The fact that many of the nests on his study area were in gull or tern colonies may have influenced the results. Hildén (1964) has shown that ducks will tolerate sparser cover than usual in order to nest among larids, an association from which they presumably receive protection against aerial predators. TABLE 2—The distribution of Lesser Scaup nests as related to the height of vegetation Nests Number (%) Height of vegetation (cm) 0-20 3( 3) 21-40 22 (39) 41-60 22 (39) 61-80 a?) > 80 5 (9) Total 56 (99) THE CANADIAN FIELD-NATURALIST Vol. 91 Lesser Scaup nests were usually well-con- cealed as compared with those of earlier nesting species. The average canopy coverage at 57 scaup nests was 35.7 + 3.6%, a figure which compares favorably with the 39.8 + 1.2% aver- age recorded for 382 Gadwall nests. The latter species has been noted for its preference for dense nesting cover (Duebbert 1966; Long 1970). The fates of 37 Lesser Scaup nests were determined. Twenty-eight (76%) were successful (i.e., at least one egg hatched), eight (22%) were destroyed by predators, and one (3%) was deserted. The nesting success for 34 insular nests was 82%. The two common predators on the study area, the Common Crow (Corvus brachy- rhynchos) and the striped skunk (Mephitis mephitis), caused most of the nest failure. Other studies also indicate that the success of island-nesting scaup is normally high (Keith 1961; Townsend 1966; Vermeer 1968; Long 1970). Under such conditions nesting success should exceed 80% and, assuming a 39% rate of renesting by unsuccessful hens (Keith 1961), more than 85% of the island-nesters should hatch successful clutches. Provision of suitable island habitat for scaup is a good management practice since the upland nests of this species are often destroyed by skunks and other predators (Keith 1961; Rogers 1964). Hatching and the Brood Period The peak of hatching was determined by backdating aged broods (Figure 3). Nearly all the successful scaup clutches hatched in July, especially during the middle two weeks of that month. Redhead clutches showed a similar but slightly earlier hatching chronology than did Lesser Scaup. The overall nesting chronology for scaup at Waterhen Marsh corresponds fairly well with that presented for central Alberta by Dwernychuk (1968). After hatching, scaup broods moved to the shallow bays protected from the wind by emergent vegetation (Figure 1). In these bays submergent plants were very sparse but aquatic invertebrates, especially chironomid larvae, were abundant (unquantified observations). The importance of such invertebrates in the diets of young scaup has been previously described (Bartonek and Hickey 1969; Bartonek and Murdy 1970; Sugden 1973). LO 40 -— SJ \J REDHEAD ( n=144>) LESSER SCAUP (n= 179) 30 20 OF CLUTCHES HATCHING % HINES: LESSER SCAUP ECOLOGY, SASKATCHEWAN 253 JULY FIGURE 3. The peaks of hatching of Lesser Scaup and Redhead broods at Waterhen Marsh. Many broods of Redheads were also observed in these areas. The preference for similar habitats by scaup and young Redhead broods, and the large populations of both species present, resulted in a great number of ducklings being concentrated into rather small areas. Asa result, mixing of broods occurred commonly. Scaup broods readily joined together since female scaup made little effort to drive away other hens or ducklings. Redhead broods were frequently deserted by their mothers, which typically showed low maternal drives. These ducklings readily joined the scaup broods with no apparent conflict resulting. In contrast to the scaup ducklings, the young Redheads remained close together and appeared to retain their identity within the mixed broods. Mixed broods of scaup were observed to come together and later separate with frequent interchange of ducklings. Female scaup showed little dis- crimination in allowing ducklings to follow them. The brood following a scaup in many cases consisted of more Redhead than scaup ducklings. Mixed broods or créches often consisted of ducklings of several age classes. The largest brood totaled more than 100 scaup ducklings and was accompanied by six hens. Typical créches contained from 15 to 40 scaup ducklings and were led by two or three hens. By early August, most of the ducklings, except those isolated from the major brood areas, were part of the créches. At this time, most of the Redhead offspring had been deserted by their mothers and 254 had also joined the large scaup broods. The mixing of broods made it difficult to determine the attrition of brood size as time went on. The average size of 78 isolated Class la broods (1 to 6 days old), believed not to have mixed with other broods, was 8.5 + 0.3 duck- lings. Because of the late-nesting habit, scaup and Redhead ducklings lagged behind other water- fowl in attaining flight. By adding the average age at first flight to the known date of hatching for the broods, the expected date of the onset of flight was calculated. Ages at first flight average about 49 days for Lesser Scaup (Gollop and Marshall 1954; Rogers 1962; Bellrose 1976) and 57 to 63 days for Redheads (Weller 1957; Smart 1965). Although Redheads showed a slightly earlier hatching chronology (Figure 3), they were later in achieving flight than were the rapidly developing scaup. By the opening of hunting season on 9 or 10 September, a maximum of 90% of the Lesser Scaup and 65% of the Redhead broods could fly. Under conditions of high early-season hunting pres- sure, local populations of these species could suffer severe losses. The Significance of Créching Behavior The formation of créches by Lesser Scaup appeared to arise because of three main conditions. First, scaup showed a relatively dense breeding population, high nesting success, and a fairly synchronized hatching period, resulting in a large number of similar-aged broods being present on the marsh at the same time. Second, there was a limited amount of suitable sheltered habitat for these broods during periods of windy weather and, conse- quently most broods assembled in the sheltered areas. The third and probably most important condition leading towards créching behavior was the high degree of tolerance, or perhaps even attraction, between different hen scaup and their broods. Scaup made no effort to drive away approaching hens with broods. Créching provides several possible advantages to the participating waterfowl species. Lack (1947) drew an analogy between this behavior in eiders and a convoy system; he suggested that in the larger broods, each female has a smaller periphery to patrol, thereby giving better THE CANADIAN FIELD-NATURALIST Vol. 91 protection against predators. Kear (1970) sug- gested that créching reduces aerial predation, as larger broods are more liable to spot approach- ing enemies and can huddle together for protection. Further evidence of this advantage is presented by Ahlen and Andersson (1970) and Gorman and Milne (1972). In the Shelduck, which in Britain undergoes an annual molt migration, the ducklings are left behind under the care of a few adults. This removes the responsibility for the care of young from most adults, which can then depart for the molting grounds (Hori 1964). Créche formation also allows more broods to use a limited amount of habitat or other restricted resource. This has been suggested for the Velvet Scoter (Melanitta fusca) by Koskimies (1955). For the Lesser Scaup, all three of these advantages could be operating. When large broods were approached by boat or canoe, two or sometimes three hens would rush at the observer tolling or feigning injury while the other hen(s) attempted to lead the brood away. Munro (1941) has reported similar behavior by scaup. Because duckling predators were not overly numerous on the study area, I did not get a chance to observe any interactions between scaup broods and their enemies. The high number of ducklings per brooding hen in the créches suggests that many hens had deserted their broods. Presumably these un- attached hens would have more time to undergo the post-nuptial molt before autumn migration. Finally, scaup broods were observed to prefer calm water areas where food was abundant. On windy days, which are frequent on the prairies, the availability of such habitat is limited. Créching allows scaup to use this habitat optimally. The mixing of Redhead ducklings with scaup broods is perhaps a further development of the parasitic habit shown by the former species in its nesting ecology. The young Redheads benefit from the protection afforded by the strong maternal drives of the scaup and the créching system. The hen Redheads can molt sooner than they otherwise might if they were still respon- sible for their ducklings. In late-nesting species such as the Redhead or Lesser Scaup, this could be of survival value as it allows earlier migration by most of the adult population. 77 Acknowledgments ] thank Maurice Porter and family of Kinistino, Saskatchewan, for the help and hospitality they extended to me while I was at Waterhen» Marsh. G. F. Ledingham kindly reviewed an earlier draft of this paper. Literature Cited Ahlen, I. and A. Andersson. 1970. Breeding ecology of an eider population on Spitsbergen. Ornis Scandinavica 1: 83-106. Bartonek, J.C. and J.J. Hickey. 1969. Food habits of Canvasbacks, Redheads, and Lesser Scaup in Manitoba. Condor 71: 280-290. Bartonek, J. C. and H. W. Murdy. 1970. Summer foods of Lesser Scaup in subarctic taiga. Arctic 23: 35-44. Bellrose, F.C. 1976. Ducks, geese, and swans of North America. Stackpole Books, Harrisburg, Pennsylvania. 544 pp. Bent, A. C. 1923. Life histories of North American wild- fowl. United States National Museum Bulletin 126. ix + 250 pp. (1962 Dover Publications reprint). Daubenmire, R. 1959. A canopy-coverage method of vegetational analysis. Northwest Science 33: 43-64. Dirschl, H. J. 1969. Foods of Lesser Scaup and Blue- winged Teal in the Saskatchewan River Delta. Journal of Wildlife Management 33: 77-87. Duebbert, H. F. 1966. Island nesting of the Gadwall in North Dakota. Wilson Bulletin 78: 12-25. Dwernychuk, L. W. 1968. Some aspects of the ecology of island-nesting waterfowl at Miquelon Lake, Alberta. M.Sc. thesis, University of Alberta, Edmonton. 155 pp. Gehrman, K.H. 1951. An ecological study of the Lesser Scaup duck (Aythya affinis Eyton) at West Medical Lake, Spokane County, Washington. M.Sc. thesis, State College of Washington, Pullman. 94 pp. Gollop, J. B. and W. H. Marshall. 1954. A guide for aging duck broods in the field. Mississippi Flyway Council Technical Section. 14 pp. (Mimeo.) Gorman, M.L. and H. Milne. 1972. Créche behavior in the Common Eider Somateria m. mollissima L. Ornis Scandinavica 3: 21-25. Hildén, O. 1964. Ecology of duck populations in the island group of Valassaaret, Gulf of Bothnia. Annales Zoologici Fennici 1: 153-279. Hochbaum, H. A. 1944. The Canvasback on a prairie marsh. American Wildlife Institute, Washington. 201 pp. Hori, J. 1964. Parental care in the Shelduck. Wildfowl Trust Annual Report 15: 100-103. Joyner, D. E. 1976. Effects of interspecific nest parasitism by Redheads and Ruddy Ducks. Journal of Wildlife Management 40: 33-38. Kear, J. 1970. The adaptive radiation of parental care in waterfowl. /n Social behaviour in birds and mammals. HINES: LESSER SCAUP ECOLOGY, SASKATCHEWAN AES) Essays on social ethology of animals and man. Edited by J. H. Crook. Academic Press, New York. pp. 357-392. Keith, L.B. 1961. A study of waterfowl ecology on small impoundments in southeastern Alberta. Wildlife Monographs, Number 6. 88 pp. Kortright, F.H. 1942. The ducks, geese and swans of North America. American Wildlife Institute, Washington. vill + 476 pp. Koskimies, J. 1955. Juvenile mortality and population balance in the Velvet Scoter (Melanitta fusca) in maritime conditions. Proceedings of the International Ornithological Conference 11: 476-479. Lack, D. 1947. The significance of clutch-size. Ibis 89: 302-352. Long, R. J. 1970. A study of nest-site selection by island- nesting anatids in central Alberta. M.Sc. thesis, University of Alberta, Edmonton. 123 pp. Munro, J. A. 1941. Studies of waterfowl in British Colum- bia, Greater Scaup duck, Lesser Scaup duck. Canadian Journal of Research, D, 19: 113-138. Rogers, J. P. 1959. Low water and Lesser Scaup repro- duction near Erickson, Manitoba. Transactions of the North American Wildlife Conference 24: 216-224. Rogers, J. P. 1962. The ecological effects of drought on reproduction of the Lesser Scaup, Aythya affinis (Eyton). Ph.D. thesis, University of Missouri, Columbia. 99 pp. Rogers, J. P. 1964. Effect of drought on reproduction of the Lesser Scaup. Journal of Wildlife Management 28: 213-222. Rogers, J. P. and L. J. Korschgen. 1966. Foods of Lesser Scaup on breeding, migration, and wintering areas. Journal of Wildlife Management 30: 258-264. Smart, G. 1965. Development and maturation of primary feathers of Redhead ducklings. Journal of Wildlife Management 29: 533-536. Sugden, L. 1973. Feeding ecology of Pintail, Gadwall, American Widgeon and Lesser Scaup ducklings. Cana- dian Wildlife Service Report Series Number 24. 45 pp. Townsend, G. H. 1966. A study of waterfowl nesting on the Saskatchewan River delta. Canadian Field-Naturalist 80: 74-88. Trauger, D. L. 1971. Population ecology of Lesser Scaup (Aythya affinis) in subarctic taiga. Ph.D. thesis, lowa State University, Ames. 121 pp. Vermeer, K. 1968. Ecological aspects of ducks nesting in high densities among larids. Wilson Bulletin 80: 78-83. Weller, M. W. 1957. Growth, weights, and plumages of the Redhead Aythya americana. Wilson Bulletin 69: 5-38. Weller, M. W. 1959. Parasitic egg-laying in the Redhead (Aythya americana) and other North American Anatidae. Ecological Monographs 29: 333-365. Weller, M.W., D.L. Trauger, and G.L. Krapu. 1969. Breeding birds of the West Mirage Islands, Great Slave Lake, NWT. Canadian Field-Naturalist 83: 344-360. Received 16 December 1975 Accepted 24 February 1977 The Genus Crangonyx (Amphipoda: Gammaridae) in the Central Connecticut River System DOUGLAS G. SMITH Museum of Zoology, University of Massachusetts, Amherst, Massachusetts 01002 Smith, Douglas G. 1977. Canadian Field-Naturalist 91(3): 256-261. The genus Crangonyx (Amphipoda: Gammaridae) in the central Connecticut River system. Abstract. The genus Crangonyx is represented in the central Connecticut River system by two poorly known species complexes, C. richmondensis and C. pseudogracilis. Examination of large series of specimens has provided ecological, distributional, and variational data for each species and has extended the zone of intergradation between C. r. richmondensis and C. r. laurentianus. A hypothesis is provided that considers glacial isolation as the factor causing differentiation in Atlantic coast populations of each species. Crangonyx r. richmondensis is considered to have re-entered New England by means of ice- contact migration while C. pseudogracilis is believed to have re-entered by way of coastal flooding. Introduction In a recent paper on North American fresh- water amphipods, Holsinger (1972) discusses taxonomic difficulties among certain widely distributed species groups, particularly Cran- gonyx pseudogracilis Bousfield and C. richmon- densis Ellis sensu lato. The problem involves insufficient distribution data for described species, and samples from given areas too small to furnish adequate information on limits of variation. The present study attempts to analyze, both systematically and zoogeographically, popula- tions of each species from the Connecticut River valley in the central New England Upland. Ecological observations have been presented for comparison with existing life-history studies and to provide information relevant to zoogeo- graphical patterns. Emphasis has been placed on examining large series of specimens representing each species in order to provide as completely as possible the relationships of central New England populations with others of the same complexes in neighboring regions. Interpreta- tion of the systematic status of Crangonyx in the Connecticut River valley relies on a proposed distributional history that accounts for the mor- phological characteristics observed in popula- tions occurring in New England. Crangonyx pseudogracilis Bousfield Between April and July 1976, 216 specimens of C. pseudogracilis were collected from 10 localities in the Connecticut River. All localities lay within Massachusetts boundaries and in- cluded Franklin, Hampshire, and Hampden Counties, or a distance of about 100 river kilometres. These were supplemented by 16 specimens taken earlier from the same river areas during May and June of 1975. All material has been placed in the Museum of Zoology, University of Massachusetts at Amherst. Also examined were nine specimens collected by E. Mills (Yale Peabody Museum Number 5473) from Axelshop Pond, Mount Carmel, New Haven County, Connecticut. Bousfield (1958) has included Atlantic coast streams within the range of this species. He also indicated that C. pseudogracilis, as a lowland river and pond species, was found in a variety of aquatic habitats. In the Connecticut River in Massachusetts, C. pseudogracilis is restricted to the river and its immediate floodplain. Intensive surveying of the entire drainage system in the state over the past three years has failed to produce specimens in any of the accessory habitats listed by Bousfield (1958). A population of C. pseudogracilis occurs, however, in the upper Quinnipiac valley west of the Connecticut valley in Connecticut (Mills 1964). Ovigerous females, occasional subadult females, and adult males mass along the shore- line in bays and oxbows from late April through mid-June. Individuals accumulate in vegetation and under rocks at the shore margin and are often found in stranded pools and rivulets formed by receding water levels. Immatures appear by the first week of June. Adult non- breeding and breeding females and males have been taken in mid-stream near bottom during late May and June and in early July. The C. pseudogracilis species complex is a 256 OTe poorly understood group discontinuously dis- tributed over greater south-central North America. The group is characterized by comb spines on the outer ramus of uropod 2 in males and weakly spined palm margins on the propods of pereopods | and 2. In the Connecticut River, individuals vary morphologically from the nominate populations examined by Bousfield (1958) in a few minor but consistent ways. Although generally resembling typical C. pseudogracilis from the Great Lakes drainages, as described by Bousfield (1958), Connecticut River specimens are distinguishable by a strongly mucronate posterolateral angle of epimeron | and the occurrence of only two to three groups of variable-length setae on the posterior margin of segments 4 and 5 on antenna 2. A quantitative assessment has shown diver- gences in some meristic characters as well. In adult females, a tendency to increase the number of apical spines on the telson lobes and to decrease spines on the posterior angle of the second gnathopod is evident. Furthermore, al- though considerable overlap exists, Connecticut River specimens are somewhat smaller than Great Lakes forms. Values and ranges for some variable charac- ters have been analyzed (Table 1). A comparison of the degree of variability within included characters shows that there is often a wide range of values for a given character and, in a few instances, that the frequency of variation is not so subtle. The presence of observed differentiation in some external features, combined with wide variability in others, may reflect evolutionary mechanisms acting upon an isolated population existing, for much of the Pleistocene Epoch, in restricted habitats in the Atlantic coastal plain (see Historical Zoogeography section). Crangonyx richmondensis Ellis During the months of March and April 1976, three populations of C. richmondensis were sampled. The collections, totaling 204 adult specimens plus additional unlisted immatures, were all taken in Massachusetts from three localities: Sunderland, Franklin County, Cran- berry Pond, 35 females, 9 males; Leverett, Franklin County, wooded vernal-autumnal SMITH: GENUS CRANGONYX IN CENTRAL CONNECTICUT 257) TABLE 1—Meristic character frequencies of Crangonyx pseudogracilis! (female, N = 113; male, N = 32) Character (female) Range Mean+ SD Propod, Gnathopod 2, posterior angle margin inner spines 3-5 35) 22 O55 outer spines 1-3 2.1+ 0.41 superior medial setae 6-10 7.8 + 0.76 Pereopod 7, posterior serrations of basis 10-16 13.4 + 0.78 Abdominal side plate 2, sub-marginal spines 2-7 43+ 1.03 Spines per telson lobe 2-6 3.7 + 0.69 Uropod 3, outer ramus, outer spine sets 3-5 317) 210290) Length (mm) males 4-5 ANS) AE LoS) females 6-10 Salt 89 ‘All specimens adults. pond, State Route 63, 13 km north of Amherst center, 59 females, 14 males; and Amherst, Hampshire County, wooded pond, State Route 9, 3 km east of center, 47 females, 40 males. All the localities listed above are small depressions in unconsolidated glacial-drift deposits filled by ground water and run-off. Each pond remains cool throughout summer. Water levels in each, except Cranberry Pond, fluctuate considerably during the year. Ovi- gerous females are present in early March and persist until at least early May. Bousfield (1958) has indicated that this species occurs in acid waters in parts of its range. The habitats of presently discussed populations maintain neutral pH within the species life zone (Keene 1968; L. Raboin, personal communication). My analysis of subspecific characters follows Bousfield’s (1958) criteria for distinguishing C.r. richmondensis and C.r. laurentianus. These include the number of apical spines per telson lobe, acuteness of posteroventral angles of the abdominal plates 2 and 3, and total length of mature specimens. Apical spines were counted on all specimens except those with damaged telsons (four individuals). Spine counts ranged from one to four per lobe, although one- and four-spined lobes were rare and have been ignored in 258 analysis. The normal arrangement was 2-2, 3-2, or 3-3. Spine counts varied both within and between populations. In Leverett and Amherst populations, 53.5% and 65.0%, respectively, had spine combinations of 2-2, indicating C. r. laurentianus, whereas 61.0% of the Sunderland population had 3-3 combinations, suggesting C. r. richmondensis. The reciprocal even com- bination occurred only in 7-17% of each; however, 3-2 combinations made up 16.0-29.5% of the remaining specimens. Total percentages of combined populations show a tendency towards C. r. /aurentianus with 48% having 2-2 combinations, the other two combinations occurring about 25% of the time. Determining the subspecific value of the second and third abdominal plates (epimera) is subjective, as the character is nonmeasurable. Figure | shows the tyical degree of develop- ment of the plate angles in all populations. Total lengths for each subspecies have been listed by Bousfield (1958) as 12 to 14 mm for females in C. r. richmondensis, and 14 to 18 mm for females and 9 to 11 mm for males in C. r. laurentianus. Mairs (1970) gives 9.9 to 13.0 mm and 6.7 to 8.5mm for females and males, respectively, for C. r. richmondensis in Maine. I observed considerable variation in inter- and intra-population length measurements. Conse- quently, subjectivity was again applied in determining overall characteristics. Sunderland and Leverett populations had lengths ranging from 13.5 to 18.5 mm and 14.0 to 19.0 mm, respectively, for females, and 10.0 to 13.0 mm and 12.5 to 18.0 mm, respectively, for males. These are typical for C.r. /aurentianus. In weg FIGURE |. Epimeral plates 1-3 of Crangonyx richmon- densis. THE CANADIAN FIELD-NATURALIST Vol. 91 20 females n-14 x-15.7 Specimens (n) Length (mm) FIGURE 2. Length curves of three populations of Crangonyx richmondensis. contrast, Amherst specimens had lengths of 12.5 to 17.5 mm and 9.0 to 12.0 mm for females and males, respectively, which is intermediate between both subspecies. Figure 2 illustrates combined values for each sex. Substantial variation occurs among local populations as demonstrated by inconsistencies in discussed characters. The Sunderland popula- tion contains apical spine counts, suggesting C.r. richmondensis, but length ranges that imply C. r. laurentianus. The reverse prevails in the Amherst population. Leverett animals generally tend toward C. r. /aurentianus in all characters. These results agree with Bell's (1971) analysis of Vermont specimens. Although he “provisionally” assigned the material to C. r. laurentianus, his examined specimens had length features of C. r. Jaurentianus and apical spine counts of C. r. richmondensis. It therefore seems unwise to consider the west-central New England forms as belonging to either subspecies. The zone of intergradation between both sub- species is then extended eastward from central New York (Holsinger 1972) to include west- central New England (Figure 3). This wide gap between seemingly good subspecies may rep- resent a non-clinal continuum resulting from mixing of two differentiated forms reinvading formerly glaciated areas (see Historical Zoo- geography section). Historical Zoogeography It is beyond the scope of this paper to recount the history of the genus Crangonyx in eastern North America. The occurrence of two seem- 1977 ingly divergent forms of two widely distributed species in northeastern North America, how- ever, warrants some discussion as to their possible origin. The effects of the Appalachian Divide on the derivation and distribution of freshwater animals is well established (Ortmann 1913; Johnson 1970). Advance of the Wisconsin ice sheet in the latter phases of the Pleistocene Epoch compressed and isolated the Atlantic coast fauna into refugia south of the ice sheet (Johnson 1970; Dadswell 1974). The position of the terminal moraine (Flint 1971) indicates that the lower coastal portions of the Susquehanna, Potomac, and Delaware River basins probably served as these refugia although the existence of refugia as far north as the exposed continental shelf areas, including Long Island Sound and Narragansett Bay (Curray 1965), has been postulated (Dadswell 1974). With the final retreat of the ice from New England during the closing phases of the epoch, avenues for dispersal became available for re- invading aquatic animals. Interdrainage pas- sages could have been negotiated either by coastal flooding along dilution zones at river mouths, by stream capture (Ross 1974) or by occupation of successional proglacial lakes, a means apparently unique, as presently known, to late-glacial events (Crocker 1957; Frey 1965; Dadswell 1974, 1975). Proglacial lake invasion, herein referred to as ice-contact migration, and stream capture would have been the only means available to animals emigrating from interior Mississippian refugia via Hudson-Champlain regions into the Connecticut valley and adjacent streams. Animals moving north from southern Atlantic glacial refugia could utilize all three methods discussed above. Due to the persistence of the major geological and topographic features of New England through the Pleistocene Epoch (Schafer and Hartshorn 1965) it is doubtful that stream capture between the Connecticut River and western systems including the Hudson and Lake Champlain drainages existed long enough, if at all, to allow faunal exchange (see Brooks and Deevey 1963, p. 150). Biological evidence in the form of non-passively distributed headwater forms is absent. The small crayfish Cambarus bartonii (Fab.) has been implicated as a typically stream-capture distributed species (Ortmann SMITH: GENUS CRANGONYX IN CENTRAL CONNECTICUT 259 1913; Crocker 1957). It commonly occurs north- ward in the upper Hudson River system (Faxon 1885; Crocker 1957), in the Champlain - St. Lawrence watersheds (Bell 1971), and has apparently entered the St. Johns River system in Maine (Faxon 1885), but has never been reported from the headwaters of any Connecti- cut River tributaries. Even the existence of an ice “tongue” that temporarily halted in the Cham- plain lowland (Lougee 1939; Schafer and Hart- shorn 1965), forming ice-marginal lakes that drained to both the Connecticut River and glacial Lake Vermont (Loveville — Fort Ann Lake succession of Coates 1976), now rep- resented by Lake Champlain, did not permit an exchange of fluvial life forms. Re-invasion of the Connecticut and adjoining eastern valleys from the interior by means of ice-contact migration apparently occurred for a few cold-water lake- dwelling fishes (Brooks and Deevey 1963), but not for cold-water lentic invertebrates (Dadswell 1974). It is suggested, therefore, that principal reoccupation of the southern New England Upland and possibly other northeastern areas by both species of Crangonyx occurred from isolated areas to the south. This is further supported by the present analysis of their mor- phology, distribution, and ecology. Moreover, the proposed methods and direction of migra- tion used by each species possibly correlates with a sequence of recolonization “waves” (Adams 1902) that included several groups of aquatic animals emigrating from southern Atlantic refugia. Crangonyx richmodensis, as previously men- tioned, occurs in water-filled kettle holes in unconsolidated stratified drift deposits typically created in ice-contact zones by meltwater streams. A definite correlation also exists between the presence of adult C. richmondensis and seasonally cold water situations (Judd 1963; Sprules 1967; Mairs 1970), implying a historical affinity of this species for cold water. Further- more, in formerly glaciated regions, this species is restricted to small lentic water bodies (Bous- field 1958). Crangonyx r. richmondensis probably left its proglacial refugia and followed closely the retreat of the ice sheet to the north by way of ice- contact migration. With abandonment and 260 subsequent isolation of ice-formed, water-filled depressions and kettle holes, C. r. richmon- densis found refuge, surviving in those areas able to maintain low water temperatures and other environmental requirements, including avoid- ance of excessive siltation and in-filling. The known localities for C. richmondensis in south- central New England lie outside the influence of heavy sedimentation typical of larger, tempo- rary glacial lakes. Crangonyx r. richmondensis eventually migrated north of the Appalachian divide. Northwesterly migration brought C. r. richmondensis in contact in valley areas with C. r. laurentianus migrating northeasterly from Mississippian refugia. This mixing resulted in interbreeding among populations of each sub- species, thus creating the observed wide band of variability that presently exists (Figure 3). During the latter stages of glacial recession C. pseudogracilis followed the same northerly route along the Atlantic coast as was suggested FIGURE 3. Proposed tracks utilized by the eastern glacial isolate Crangonyx richmondensis richmodensis during postglacial dispersal. Shaded area depicts zone of intergradation of both species. Map key: D = Delaware River, H = Hudson River, C = Con- necticut River, M = Mohawk River, LO = Lake Ontario, LC = Lake Champlain; dashed line = Ter- minal moraine of Pleistocene glaciation, solid line = Appalachian Divide, arrows = direction of dis- persal, 18000 B.P. shoreline during maximum glaciation follows Curray (1965). THE CANADIAN FIELD-NATURALIST Vol. 91 FIGURE 4. Proposed track utilized by the eastern glacial isolate Crangonyx pseudogracilis during postglacial dispersal. See Figure 3 for map key. Shaded area depicts known present distribution of the species in study area. for the two crayfish Orconectes limosus (Raf.) and Procambarus a. acutus (Harland) by Ortmann (1906) and Crocker (1957) (Figure 4). Dispersal probably was controlled by flooding processes in coastal zones correlated with a rising sea-level. A review of the ecology of C. pseudogracilis in this study and in Bousfield (1958) shows that this species is a lowland river and pond inhabitant. Observations discussed earlier indicate an annual shoreward migration in spring. Animals massing in shallows near river mouths during spring floods could move between drainage basins. Adventives of C. pseudogracilis most likely entered the southern Connecticut coastal plain during glacial Lake Hitchcock times, in the period following the Middletown re-advance about 13000 B.P. Between the time of glacial lake formation and final draining about 10700 B.P. (Schafer and Hartshorn 1965; Hartshorn 1969) C. pseudogracilis spread north- ward through the temporarily common Quinnipiac-Farmington valley lake-drainage system (Lougee 1938) as indicated by the presence of a relict population (E. Mills collection). Further eastward, the species entered the Connecticut Valley ‘and passed northward through the New Britain spillway into Lake Hitchcock, extending at least to the Turners Falls bedrock barrier. Crustal upwarping, following lake drainage, increased the incipient 1977 rivers gradient downward to the south. This caused increased downcutting and eventual draining of former lake basin areas. Crangonyx pseudogracilis, having just occupied the lake, possibly was trapped by the sudden physio- graphic changes brought about by upwarping and as yet, has not left the immediate river valley. Acknowledgment This paper was published through assistance from the Guy Chester Crampton Research Fund, University of Massachusetts. Literature Cited Adams, C. C. 1902. Postglacial origin and migrations of the life of the northeastern United States. Journal of Geography 1: 352-357. Bell, R.T. 1971. Handbook of the Malacostraca of Vermont. Privately published, Burlington, Vermont. 65 pp. Bousfield, E. L. 1958. Fresh-water amphipod crustaceans of glaciated North America. Canadian Field-Naturalist 72(2): 55-113. Brooks, J. L. and E.S. Deevey. 1963. New England. /n Limnology of North America. Edited by D.G. Frey. University of Wisconsin Press, Madison, Wisconsin. pp. 117-162. Coates, D. R. 1976. Quaternary stratigraphy of New York and Pennsylvania. /n Quaternary stratigraphy of North America. Edited by W. C. Mahaney. Halsted Press, New York. pp. 65-90. Crocker, D. W. 1957. The crayfishes of New York state. New York State Museum and Science Service Bulletin 355. 97 pp. Curray, J. R. 1965. Late Quaternary history, continental shelves of the United States. Jn The Quaternary of the United States. Edited by H. E. Wright, Jr. and D. G. Frey. Princeton University Press, Princeton, New Jersey. pp. 723-735. Dadswell, M.J. 1974. Distribution, ecology and _ post- glacial dispersal of certain crustaceans and fishes in eastern North America. National Museum of Canada, Publications in Zoology 11. 110 pp. Dadswell, M. J. 1975. Further new localities for certain coldwater fishes in eastern Ontario and western Quebec. Canadian Field-Naturalist 89(4): 447-450. Faxon, W. 1885. A revision of the Astacidae. Part I. The genera Cambarus and Astacus. Memoirs of the Museum of Comparative Zoology 10: 1-186. Flint, R. F. 1971. Glacial and Quaternary geology. John Wiley, New York. 892 pp. SMITH: GENUS CRANGONYX IN CENTRAL CONNECTICUT 26] Frey, D. G. 1965. Other invertebrates—An essay on bio- geography. /n The Quaternary of the United States. Edited by H. E. Wright, Jr. and D. G. Frey. Princeton University Press, Princeton, New Jersey. pp. 613-631. Hartshorn, J. H. 1969. Geography and geology of glacial Lake Hitchcock. /m An introduction to the archeology and history of the Connecticut Valley Indian. Edited by W. R. Young. Springfield Museum of Sciences Publications, New Series 1(1): 19-28. Holsinger, J. R. 1972. The freshwater amphipod crusta- ceans (Gammaridae) of North America. /n Biota of freshwater ecosystems. United States Environmental Protection Agency. Identification Manual 5. 89 pp. Johnson, R. I. 1970. The systematics and zoogeography of the Unionidae of the southern Atlantic slope region. Bulletin of the Museum of Comparative Zoology 140(6): 263-449. Judd, W. W. 1963. Studies of the Byron Bog in south- western Ontario. XVI. Observations on the life cycles of two species of Crangonyx. National Museum of Canada, Natural History Papers 20: 1-9. Keene, C. I. 1968. Some ecological parameters of Cran- berry Pend, Sunderland, Massachusetts. M.Sc. thesis, University of Massachusetts, Amherst. 62 pp. Lougee, R.J. 1938. Physiography of the Quinnipiac- Farmington Lowland in Connecticut. Colby Monographs 7. 64 pp. Lougee, R. J. 1939. Geology of the Connecticut water- shed. New Hampshire Fish and Game Department, Biological Survey of the Connecticut Watershed Depart- ment 4: 131-149. Mairs, D.F. 1970. Observations on the life history of Crangonyx richmondensis richmondensis Ellis. American Midland Naturalist 83: 315-318. Mills, E. L. 1964. Noteworthy Amphipodain the collection of the Yale Peabody Museum. Postilla 79. 41 pp. Ortmann, A. E. 1906. The crawfishes of the State of Pennsylvania. Memoirs of the Carnegie Museum 2(10): 343-523. Ortmann, A.E. 1913. The Alleghenian Divide and its influence upon the freshwater fauna. Proceedings of the American Philosophical Society 52: 287-390. Ross, H. H. 1974. Biological systematics. Addison-Wesley, Reading, Massachusetts. 345 pp. Schafer, J. P. and J. H. Hartshorn. 1965. The Quaternary of New England. /n The Quaternary of the United States, a review volume for the VIIth Congress of the International Association for Quaternary Research. Edited by H.E. Wright and D.R. Frey. Princeton University Press, Princeton, New Jersey. pp. 113-128. Sprules, W. G. 1967. The life cycle of Crangonyx richmon- densis laurentianus Bousfield. Canadian Journal of Zoology 45: 877-884. Received 6 August 1976 Accepted 8 April 1977 Pteridophytes of the Regional Municipality of Waterloo, Ontario CRAIG A. CAMPBELL! and DONALD M. BRITTON? 1421 King Street North, Waterloo, Ontario N2J 3Z4 2Department of Botany and Genetics, University of Guelph, Guelph, Ontario NIG 2W1 Campbell, Craig A. and Donald M. Britton. 1977. Pteridophytes of the Regional Municipality of Waterloo, Ontario. Canadian Field-Naturalist 91(3): 262-268. Abstract. Sixty-eight taxa of pteridophytes are listed for Waterloo Region, consisting of 51 species and 17 hybrids, varieties, and forms. Eleven species and 13 hybrids, varieties, and forms are new additions to the published flora, our knowledge of which was based on the pioneering work of William Herriott around 1900, and F. H. Montgomery in the 1940s. The status of rare or scarce pteridophytes in the region is discussed, and nine species that could not be found after 10 years of field work but might be expected to occur in the region, are enumerated. Key Words: pteridophytes, Waterloo region Ontario, additions to flora. The earliest known check list of pteridophytes of Waterloo County (now the Regional Munici- pality of Waterloo) was the unpublished one of William Herriott of Galt (now Cambridge). This list is undated, but an annotation indicates a date of circa 1926. Herriott listed 29 taxa of ferns only. Montgomery (1945) with reference to Herriott’s collections and his own field work, wrote “A Botanical Survey of Waterloo County.” Forty-three taxa of pteridophytes, 32 of these ferns, were recorded in it. For almost 25 years after this, no systematic work was done on the Waterloo County flora. From 1959 on, Britton has been studying the biosystematics of Dryopteris, making col- lections in the Waterloo region. Since 1967, Campbell has been reinventorying the flora of the area, an imperative undertaking because of rapid urbanization. Persons associated with the University of Waterloo and Wilfrid Laurier University have also been engaged in such a resurvey. In addition to extensive field searches for pteridophytes in the Waterloo region, we have examined specimens in the following herbaria: CAN, DAO, HAM, LKHD, NFO, OAC, QK, TRT, TRTE, UWO, WAT, WIND, WLU. Literature and specimens from surrounding areas within the Great Lakes basin were also examined for comparative purposes. In general, we have followed the nomen- clature of Wherry (1961). Where these names differ from those used by Fernald (1950) and Montgomery (1945), the synonomy is given. To indicate status within the region, we have used the following code: | to 2 locales, rare; 3 to 5, scarce; 6 to 10, occasional; 11 to 20, frequent; over 20, common. In addition, an indication of abundance at some sites has been given. With rare species, we have tended to emphasize their history in the region. We endeavored to check all stations of rarer taxa, as well as all reports of them. Repre- sentative collections were made of most com- mon taxa. Collections made from 1967 to 1976 by Campbell and associates are largely in the herbarium of Wilfrid Laurier University (WLU). Duplicates of some have been deposited in the National Museum Herbarium, Ottawa (CAN). The classification of the Pteridophyta is in a state of flux at this time. The system we have adopted is that of Crabbe et al. (1975). This system is noteworthy in placing many of our ferns in the family Aspleniaceae and only one (Polypodium) in the Polypodiaceae. For sim- plicity and clarity, we have grouped the numerous Dryopteris hybrids into one section, whereas three Equisetum hybrids are integrated into the Equisetum sequence. We have ignored trivial ecological or mutant forms and varieties, but have attempted to encompass those varieties, forms, and hybrids which are considered important by the leading pteridologists of today. 262 ST Lycopodiaceae *Lycopodium annotinum L. Scarce; recorded from five stations, numerous at one, a white spruce bog at Kossuth, Campbell and Lamb 70-7 (WLU). Others under hemlock in old mixed forests at Erbsville, Sunfish Lake, and Bamberg (Wellwood 2018, Campbell and Pratt 71-71, Campbell and Diebolt 75- 93 [all WLU]). Also at Spongy Lake (Lamb s. n., WLU). Lycopodium clavatum L. Occasional. Collected at nine locations, but sparse; e.g., spruce bog, Kossuth, Campbell and Lamb 70-6 (WLU), Idlewood Park, Kitchener, Campbell and R. Britton 75-15 (WLU), Hidden Valley Road, Kitchener, Campbell et al. 71-74 (CAN). Only earlier collections from Galt “near old sawmill” along Mill Creek, No. 228 and Gibson’s Woods, North Dumfries Township, No. 229 ( Herriott in OAC). Lycopodium complanatum L. var. flabelliforme Fernald (in Montgomery (1945) as Lycopodium complanatum L.). Scarce; collected at four stations, and sparse. Montgomery (1944) reported it only at Orr’s Lake, North Dumfries Township, concession XI, lot 17, Herriott Acc. No. 226 (OAC); mixed woods 1 mi (1.6km) south of Crosshill, Campbell (8); Idlewood Park, Kitchener, Campbell and Lamb 70-8; Dryden Tract, North Dumfries Township, concession IX, lot 21, Campbell and Diebolt 75-86 (all WLU). At the last site, a large infertile colony apparently adventive into 15-year-old pine plantation on edge of hardwoods. *Lycopodium inundatum L. var. inundatum. Rare, as it is elsewhere south of the Precambrian Shield in Ontario. Open, highly acidic, sphagnum bogs; in small, sparsely-vegetated depressions; Herriott’s Bog, 3 mi (4.8 km) south of Galt (this location also known as Oliver’s Pond, in North Dumfries Township, concession VIII-IX, lot 6), Montgomery 1561 (OAC) and also from here, Britton and Campbell 75-157(2) (WLU). Sight record: Grass Lake (Paris Cranberry Bog), North Dumfries Township, concession VII, lots 17, 18, R. MacLaren in 1975. Lycopodium lucidulum Michaux. Frequent. Lycopodium obscurum L. Occasional in dryish mixed woods. The typical variety and var. *dendroideum D. C. Eaton are both represented in the collections, e.g., Wellwood 2070 (WLU) from Spongy Lake, and Wellwood 622 (WLU) from Erbsville, respectively. As Gleason (1952) notes, these two varieties seem to intergrade. *Taxon new to the published flora of the Regional Municipality of Waterloo. CAMPBELL AND BRITTON: PTERIDOPHYTES, WATERLOO, ONTARIO Selaginellaceae Selaginella apoda(.) Fernald. Occasional. Collected at seven localities; damp marly river, stream, and lake banks; also limy meadow and mossy edge of cedar swamp. Herriott, west side [of Grand River below Galt], No. 224 (OAC), Sunfish Lake, open meadow, Britton and Peterson 1452 (OAC), Blair Swamp, Beasley’s old survey, lot 3, bank of rill, Campbell et al. 72-46 (CAN) are representative collections. Equisetaceae Equisetum arvense L. Common. Some collections have been identified as var. *boreale Ruprecht: Campbell 72-38 (WLU), 73-30 (CAN), for example; these were from wet, forested localities. Equisetum fluviatile L. (formerly Equisetum limosum L.). Occasional, in bogs, lakes, marshes, and along waterways and ditches; sometimes numerous. Equisetum hyemale L. (Equisetum prealtum Raf. in Montgomery (1945)). Frequent. We have only var. affine (Engelmann) A. A. Eaton. * Equisetum hyemale var. affine X Equisetum laeviga- tum (= Equisetum X ferrissii Clute). Rare. Quite numerous at the one location on damp sand floor of gravel pits, along a ridge, Hidden Valley Road, Kitchener, Campbell et al. 74-316 (WLU), Britton 3343 (OAC). These collections were determined by R. L. Hauke. *Equisetum hyemale var. affine X Equisetum vari- egatum ( = Equisetum X trachydon A. Braun). Rare. Few at one location in gravel pit on Bird Ridge, Hidden Valley Road, Kitchener, August 1975, Anderson s.n. (OAC). *Equisetum laevigatum A. Braun. Scarce. Sparse to numerous at three locations in the region; a small seepage area at source of Patterson Creek, Beake Pond (= Taylor Lake) south of Cambridge, Camp- bell et al. 74-303 (WLU), Britton 3335 (OAC); damp floors of abandoned gravel pits. Hidden Valley (Bird Ridge) Road, Kitchener, Campbell et al. 74-176 (WLU), Britton 3341 (OAC). Determined by R. L. Hauke. Railway ditch. Galt [Cambridge], Mont- gomery 15-39(TRT), reported by Montgomery (1945) as Equisetum prealtum Raf. and revised by Taylor and Boivin to Equisetum laevigatum. Typically a western species. *Equisetum laevigatum X Equisetum variegatum (= Equisetum X nelsonii (A. A. Eaton) Schaffner). Rare. Beake Pond, Britton et al. 3973 (OAC), determined by R. L. Hauke; along Blair-Galt Road, Campbell et al. 75-120(2) (WLU). * Equisetum pratense Ehrhart. Scarce. Occurs at three locations in small patches surrounded by Equisetum arvense; sites are moist or springy shaded areas, in rich 264 organic soil or sand; Homer Watson Park, Kitchener, Campbell and Schaefer 74-107 (WLU, CAN), Campbell and Donaldson 74-123 (WLU), and Strasburg woods, Kitchener, Campbell and Diebolt 75-79(4) (WLU); Spongy Lake, McIntosh et al., 29 September 1976 (OAC). More common northward in Ontario. Equisetum scirpoides Michaux. Occasional. Equisetum sylvaticum L. Occasional. Apparently localized in cedar swamps and bog margins and sometimes climax mixed forests near water; abundant only at Paradise Lake. Some sites are Roseville Swamp (North Dumfries Township, concession IX, lot 27), Campbell 69 s. n. (WLU); Idlewood Park hydro corridor, Kitchener, Campbell and R. Britton 72-73 (WLU), St. Agatha (Wilmot Township, concession | of Block B, lot 3), Campbell and Donaldson 74-13 (WLU). Varieties and/or forms were not considered. Equisetum variegatum Schleicher. Occasional, but localized in limy regions along rivers and streams; also in marl meadows; abandoned gravel pits where probably adventive. A luxuriant colony occurs at West Montrose, Highway 86 bridge, northeast bank of Grand River, Campbell and Bald 73-40 (CAN). Montgomery (1944) recorded only one station: Hogg’s swamp at the riverside, Galt, 15 June 1894, Herriott (OAC), now a landfill site. Ophioglossaceae Ophioglossum vulgatum L. var. pseudopodum (Blake) Farwell. Rare; collected only at Sunfish Lake near Erbsville in 1940, Montgomery 445 (OAC). Repeated searches by the authors for it here failed. Botrychium dissectum Sprengel. Occasional; col- lected at eight locations, few in number in hardwoods (often beech-maple) or rich mixed forests. Forma *dissectum Sprengel, Ridgeway Drive, Cambridge, Campbell et al. 72-47 (CAN) and Bamberg, Camp- bell and Diebolt 75-89 (WLU). Forma obliquum (Muhlenberg) Fernald represented by Freeport, Montgomery s. n. (OAC Acc. No. 1619) in 1947 and Crosshill, Campbell and England s.n.(WLU Acc. No. 4719). One intermediate between these two formae from Crosshill site (WLU Acc. No. 47-20). Variety oneidense (Gilbert) Farwell has been found only twice: once at Shoemaker’s Woods, Kitchener, Montgomery 446 (OAC) (determined by W.H. Wagner, Jr. as Botrychium oneidense [Gilbert] House), and a sight record at Dickson Wilderness, by Campbell and Britton, in October 1974, in same habitat as multifidum and dissectum. * Botrychium matricariaefolium A. Braun. Rare; one collection from open knoll in yellow birch — red maple — white cedar swamp near Bamberg, Wellesley THE CANADIAN FIELD-NATURALIST Vol. 91 Township, concession VI, Campbell and Diebolt 75- 88 (CAN, WLU). * Botrychium multifidum (Gmelin) Ruprecht. Scarce. Definitely known at four localities only: Strasburg, Kitchener, Campbell and Britton 70-168 (WLU) and Campbell and Bald 73-11 (CAN); Salisbury Drive (west of Victoria Park), Cambridge (Galt), Campbell and Campbell 75-76 (WLU); also seen at Dickson Wilderness, Campbell and Britton, November 1975. On edges of sandy, deciduous white pine woods, plants not robust. Collected by Montgomery, 1939, as Botrychium obliquum (Montgomery 446, HAM) at Kitchener. Botrychium simplex E. Hitchcock. Rare. One location, Dickson’s Woods, Galt, open grassy place, Prescott in 1889 (OAC Acc. No. 208) and thicket in edge of swamp, Salisbury Road west of Dickson’s Woods, Cambridge (Galt), Campbell (3)(WLU). Ona sheet at QK (Acc. No. 00519), Herriott, Galt in 1904, there are varieties simplex, *laxifolium Clausen and *tenebrosum Clausen as determined by Britton and Campbell 1973. Botrychium virginianum Swartz. Common. Osmundaceae Osmundae cinnamomea L. Frequent. * Osmunda claytoniana L. Herriott (without date) lists this species as “frequent”. No Herriott specimens have, however, been located. Occasional, edges of swamps and low woods and bases of slopes, e.g., Paradise Lake, Sudden Tract (Causeway Swamp), northwest of Galt “Footbridge,” Dryden Tract (Alps Road); Dickson Wilderness, Brachton and University of Waterloo campus (seven locations). Osmunda regalis L. var. spectabilis (Willdenow) Gray. Frequent. Adiantaceae Pellaea glabella Mettenius var. glabella. Rare. Known only from dolomitic cliffs on both banks of the Grand River between Galt and Preston (Cambridge): abundant on some cliffs. Collected here as early as 1893 by Herriott and intermittently to the present. Early collections were identified as Pellaea atro- purpurea (L.) Link, a triploid, but are referable to Pellaea glabella, a tetraploid which is glabrous. A recent voucher: Campbell and Reznicek 71-95 (OAC). Adiantium pedatum L. Frequent. Polypodiaceae Polypodium virginianum L. Scarce. Collected at three sites. On dolomitic outcrops at the rifle range north of Galt, Herriott, August 13, 1892 (OAC) and Wilke’s estate (Cruickston Farm), Campbell and Schaefer 70- 173 (WLU); and on tree roots, Northeast Woolwich ISTH Swamp, Campbell and Lamb 24 May 1969 (WLU). Sight records: Doon Pioneer Village (Campbell circa 1964); Grass lake (Paris Cranberry Bog), Macdonald, International Biological Program Survey, on logs and wooded banks, respectively; also, at Josephburg and Erbsville (Diebolt; Lamb 15, 1976, large colony on root mass). Our plants are sexual tetraploids ( = 74) Dennstaedtiaceae Dennstaedtia punctilobula (Michaux) Moore. Scarce, collected at three localities in mixed and deciduous woods. Four mi (6.4 km) southwest of Kitchener, Montgomery 774 (OAC) and 14 mi (2.4 km) below #7 Highway near Petersburg, Mont- gomery 774 (TRT), are presumably the same site; Doon Pioneer Village, Kitchener, Campbell and Bald 73-14 (CAN); Schaefer’s Woods, Erbsville, Campbell 75-135 (WLU). Pteridium aquilinum (L.) Kuhn var. /atiusculum (Desvaux) Underwood ex Heller. Common. Thelypteridaceae Thelypteris noveboracensis (L.) Nieuwland (as Dryop- tersis in Gray’s Manual, Fernald (1950)). Occasional, collected at eight localities: e.g., Branchton West, Herriott, 20 September, 1902 (OAC), Pioneer Village Woods, Campbell, 11 June, 1964 (WLU), Erbsville, Wellwood 312 (WLU). Thelypteris palustris Schott var. pubescens Fernald (as Dryopteris thelypteris (L.) Gray in Gray’s Manual (Fernald 1950)). Common. * Phegopteris connectilis (Michaux) Watt (as Dryo- pteris phegopteris (L.) Christensen in Gray’s Manual, Fernald (1950)). Scarce, collected at four localities, in or along swamps under beech or cedar, few; Wilke’s estate, Herriott (OAC 1939), Paradise Lake, Mont- gomery 1029 (OAC), Roseville Swamp, Campbell s.n. 27 July 1969 (WLU), and Schaefer’s Woods, Erbsville, Campbell et al. 75-123 (WLU). Aspleniaceae * Asplenium platyneuron (L.) Oakes. Rare, in Dryden Tract (Alps Road), North Dumfries Township; small widely-scattered colony growing on moist sand in pine reforestation; photographed here in September 1975; substantiating earlier report of Diebolt and Donald- son for 8 July 1975 in the Waterloo Region Annotated Plant Species List, 1976, Man-Environment Studies, University of Waterloo. A second colony was found in the Sandy Hills Tract, (North) Woolwich Township: eight young plants on shady, mossy ridge of sandy loam in maturing red pine and spruce plantation, found by G. Francis, 21 May 1977, Campbell 77-41 (CAN, WLU). Asplenium trichomanes L. Rare, well-known from CAMPBELL AND BRITTON: PTERIDOPHYTES, WATERLOO, ONTARIO 265 exposed dolomitic outcrops of Cruickston Park Farm (Wilke’s) along Grand River, and opposite in City of Cambridge (Preston); Herriott, 1895 (OAC, 170), Montgomery and Campbell s. n., 2 November 1968 (WLU). Our collections are tetraploid ( = 72). Camptosorus rhizophyllus (L.) Link. Known from the collections of Herriott, in 1893 (OAC) and McGill (without date and detailed locality), QK. Considered extinct by Montgomery (1945). Herriott’s locality was “the rifle-range, Galt,” which now is sparsely shaded and appears too dry to support this species. Thorough searches have been made for it. Matteuccia pensylvanica (Willdenow) Morton (as Pteretis nodulosa (Michaux) Nieuwland in Mont- gomery (1945)). Common. Onoclea sensibilis L. Common. Athyrium filix-femina (L.) Roth (as Athyrium ang- ustum [Willdenow] Pres] in Montgomery (1945)). * Athyrium pycnocarpon (Sprengel) Tidestrom. Scarce, collected at five localities where it occurs sparsely in rich moist seepage areas in hardwoods: Wilke’s estate, J. Kerr, 1904 (OAC); | mi (1.6 km) south of Galt, Cruise 8602, 8 September 1956 (TRT, OAC, UWO); Waterloo (Rummelhardt), Kott and Kott, 15 August 1972 (WLU); Bamberg, Diebolt and Campbell 75-87 (2), 14 September 1975 (WLU, CAN); Schaefer’s Woods, Erbsville, Campbell 75-136, 5 October 1975 (CAN, WLU); and McGill (QK) without date or detailed locality. Also, sight records are known for the St. Agatha, West Montrose, and Kitchener areas (Brown 1975; MacDonald 1970; Shantz circa 1962). Athyrium thelypterioides (Michaux) Desvaux. Oc- casional. In rich, moist deciduous woods on seepage banks and in glades: for example, Galt, Herriott, 10 July 1894 (OAC); Sugar Bush Park, Waterloo, Campbell s.n., June 1968 (WLU); north of West Montrose, Britton et al. s. n., 26 July 1972 (OAC). Gymnocarpium dryopteris (L.) Newman (Dryopteris linnaeana Christensen in Montgomery (1945) and Dryopteris disjuncta Morton in Fernald (1950)). Occasional in cedar swamps and low woods under conifers: for example, Paradise Lake, Britton et al. 2658, 26 July 1972 (OAC); Preston, Lewis, s. n., 22 June 1936 (TRT). Cystopteris bulbifera (L.) Bernhardi. Frequent. Cystopteris fragilis (L.) Bernhardi. Occasional. Usu- ally on moist, shady banks or sandy ridges: for example, Spongy Lake, Wellwood 2104, (WLU); Alps Woods, Wellwood 731(WLU); Ayr Road, Adams 249 (WAT). All specimens we have seen for the region are referable to variety mackayi Lawson. Adams 249, although very large and growing on soil, has spores conforming in size to tetraploid C. fragilis, not to 266 those of C. protrusa Blasdell, a diploid. Polystichum acrostichoides (Michaux) Schott. Com- mon. Dryopteris clintoniana (D. C. Eaton) Dowell ( Dryop- teris cristata var. clintoniana Underwood in Mont- gomery (1945) and Fernald (1950)). Frequent, some- times numerous, in rich, springy ground or on rotten logs and bases of dead trees. Found in diverse habitats from deciduous woods to glades in white spruce-cedar swamps. A cytological voucher (n=123) from Altrieve Lake, North Dumfries Township, Britton B534 (OAC). Dryopteris cristata (L.) A. Gray. Occasional. A cyto- logical voucher from Altrieve Lake bog, Britton 528 (OAC) (n = 82). Dryopteris goldiana (Hooker) A. Gray. Rare, known only from two localities: one extant colony: “Rum- melhardt” near Erbsville, small patch in rich de- ciduous woodlot near intermittent stream, Wellwood 2129 (WLU). The stand in the swamp at back of Dickson’s Woods (Victoria Park), Cambridge (Galt), Herriott in 1902 (OAC) has not been relocated despite repeated search. A sheet at QK (Acc. No. 50928) collected by D. H. McGill without date or locality except “Waterloo County” is this species. Dryopteris intermedia (Willdenow) A. Gray (Dryop- teris spinulosa var. intermedia Underwood in Mont- gomery (1945) and Fernald (1950)). Frequent, in mixed or rich deciduous woods, for example, Glas- gow Woods, golf course, Kitchener, Campbell 73-47B and “Rummelhardt,” Waterloo, Campbell and R. Britton 72-22, and Schaefer's Woods, Erbsville, Campbell 75-137 (all WLU). Dryopteris marginalis (L.) A. Gray. Occasional. Dryopteris spinulosa (O. F. Mueller) Watt (Dryop- teris carthusiana (Villars) H. P. Fuchs). Montgomery (1944) and the Waterloo Region Annotated Plant Species List (1976) provide numerous records. Mont- gomery (1944) cites a specimen of Herriott(OAC 210) as Dryopteris spinulosa var. americana (Fischer) Fernald. This taxon is now considered as comprising two sexual species, Dryopteris assimilis S. Walker, a diploid, presently not known from nearer than eastern Lake Superior, and Dryopteris campyloptera (Kunze) Clarkson, essentially an Appalachian species still unconfirmed for Ontario. The Herriott sheet (OAC 210) is comprised of two taxa: one frond of Dryopteris intermedia and a portion of a frond Dryopteris X trip- loidea Wherry. Hybrids Dryopteris clintoniana X cristata. Apparently rare, collected only at Altrieve Lake, with parents, low wet swamp, Britton 532 and 533 (OAC), the latter a cyto- THE CANADIAN FIELD-NATURALIST Vol. 91 logical voucher (82 II, 41 I) (Widen and Britton 1971). *Dryopteris clintoniana X intermedia (Dryopteris X dowellii Wherry). Occasional, collected at six sites in company of parents, e.g., springy ground and swamp edges and glades, Strasburg Road, Kitchener, Camp- bell and R. Britton 72-5 (WLU), Homer Watson Park, Kitchener, Campbell and R. Britton s.n. (WLU 9596), Beake Pond, R. Britton 6 September 1972 (WLU). *Dryopteris clintoniana X marginalis (Dryopteris X burgessii Boivin). Scarce, collected in Roseville Swamp in hemlock-yellow birch stand, Britton 2518 (OAC), Wilmot Center, Wilmot Township, boggy swamp edge, Campbell 72-27 (CAN, OAC) and Homer Watson Park, Kitchener, Campbell and Schaefer 75-46 (OAC). Rarely collected (Wherry 1961). Dryopteris cristata X intermedia (Dryopteris X boot- tii [Tuckerman] Underwood). Occasional with par- ents in swamps and moist woods: Altrieve Lake, Britton B536 (OAC), a cytological voucher (2n = 123), Northeast Woolwich Swamp, Britton and Campbell 2239 (OAC), Beake Pond, Campbell et al. 74-301 (OAC), Homer Watson Park, Kitchener, Campbell and Schaefer 74-249a (WLU) are representative col- lections (Widén and Britton 1971). * Dryopteris cristata X marginalis (Dryopteris X slos- sonae Wherry). Rare, collected only once, in muck among cedars, Doon Pioneer Village, Kitchener, Campbell and Bald 73-13 (WLU). Dryopteris cristata X spinulosa (Dryopteris X uligi- nosa Druce). Rare, collected twice, in swamps with parents, Beake Pond, R. Britton 6 September 1972 (WLU) and Altrieve Lake, Britton 535 (OAC) (Widén and Britton 1971). *Dryopteris intermedia X spinulosa (Dryopteris X triploidea Wherry). Frequent. *Dryopteris marginalis X spinulosa (Dryopteris X pittsfordensis Slosson). Rare, collected once, to north of Dickson Wilderness, North Dumfries Township, Campbell s. n., 4 October 1970 (WLU). Blechnaceae * Woodwardia virginica(L.) J. E.Smith. Rare, known only from the remnants of a large peat bog (now Idlewood Park), Kitchener: R. Britton and Campbell 2-34, 15 August 1972 (WLU, CAN, OAC). Dis- covered here by V. Shantz circa 1971. Very scattered across southern Ontario, limited to certain bogs; mapped by Cody (1963). (A report from Beverly Swamp in the Waterloo Region Annotated Plant Species List (1976) may refer to Wentworth Region; there is no known specimen for the Waterloo portion of the Swamp.) LOTT Exclusions Equisetum palustre L. Although reported in the Waterloo Region Annotated Plant Species List (1976), from three localities in the region, no specimens can be located. The species should be present. Phegopteris hexagonoptera (Michaux) Feée. This species has not been found in the region, although it is well known from Pinehurst Park and Spottiswood Lake just south of the regional boundary in Brant County. The specimen at QK (Herriott, 17 September 1904) is a duplicate of Herriott’s sheet in OAC, but is missing the detailed locality datum which was “Spottiswood Lake.” The species should be present in the region. Polystichum braunii (Spenner) Fée. A “Waterloo County” collection by McGill, “July” (1913 added by someone else) is in QK (Acc. No. 50904). This northern fern has its nearest known definite locality at Nestleton Station, Durham County (Taylor 1934); its occurrence in Waterloo region seems unlikely. Polystichum lonchitis (L.) Roth. Reported by Montgomery (1944) as having been collected by Herriott at the “footbridge below Galt,” 13 July 1892; the specimen has not been seen by the authors, and its whereabouts is not known. Very little suitable habitat seems to exist at this site today. Discussion This paper brings to 68 the taxa of pterid- ophytes definitely known from the Waterloo region. Eleven of these are hybrids (three Equisetum, eight Dryopteris) and six are often considered as varieties, or well-marked forms, hence 51 are species. (Another four taxa are here excluded from the regional flora.) Of these 51 species, 11(*) are herein reported for the first time for the Waterloo region; in addition, 13 varieties, forms, and hybrids are reported for the region for the first time. (See footnote on page 33.) This list of pteridophytes has more taxa than lists for neighboring counties do. Although no complete lists as yet exist for Perth, Oxford, Brant, or Wentworth Counties, Britton has records for 47 species from Wellington County and Cruise (1969) lists 45 native species for Norfolk County. In the Waterloo region, 10 species of club mosses and ferns are rare and knownat only one or two Stations, whereas only seven are common. Seven hybrids are also considered rare at CAMPBELL AND BRITTON: PTERIDOPHYTES, WATERLOO, ONTARIO 267 present, but may be overlooked at other sites. One species, the Walking Fern (Camptosorus rhizophyllus) appears to be extinct in the region; also, the Adder’s Tongue (Ophioglossum vul- gatum) has not been recorded recently. Recent designation (1976) of environmentally sensitive areas within the region may provide some protection for a number of stations of rare pteridophytes. One of the plants listed for Ontario as rare by Argus and White in a mimeo list from the National Museum is the Narrow- leaved Spleenwort (Athyrium pycnocarpon), it is now recorded from eight locations in the Waterloo region but very few living vigorous clones exist. A number of the species, herein listed as rare regionally, might also be considered endangered. Although Dryopteris goldiana has decreased within the region, and the holly ferns (Polystichum lochitis and Polystichum braunii), if they actually occurred here, may be extinct, several pteridophytes may be increasing. Abandoned gravel pits seem to provide habitats for new stations of Equisetum variegatum and Equisetum laevigatum. Possibly the grazing of many woodlots as well as walking trails through them has provided suitable ecological niches for colonies of botrychiums. Also, Britton believes that the Ebony Spleenwort, Asplenium platy- neuron is increasing in southern Ontario, which may account for our recent addition of it to the Waterloo flora. Dryopteris clintoniana and Dryopteris hybrids undoubtedly have not in- creased, but because of particular attention paid to this genus, have been found more often. Despite some searching for them, nine species which are to be expected in the region have not been found yet. In addition to the two in the excluded list, they are: Lycopodium trista- chyum, Selaginella rupestris (both on sand), Botrychium lanceolatum, and Botrychium ter- natum, Cryptogramma Stelleri, Asplenium viri- de, and Pellaea atropurpurea (which all might conceivably be on the cliffs at Cruickston Farm). The hybrid horsetail Equisetum X litorale (Equisetum fluviatile X Equisetum arvense) likely occurs in the Waterloo region, as well as some other hybrid pteridophytes, especially in Dryopteris. The diversity of habitats within the Regional Municipality of Waterloo surely contributes to 268 the relatively rich (for southwestern Ontario) pteridophyte flora. Numerous swamps and bogs, as well as many rich woods, provide a range of acidic to neutral soils. Rock outcrops are few, as are sandy tracts, but both contribute a few notable species such as Equisetum laevi- gatum and Asplenium trichomanes. Closely associated with limestone (dolomitic) rock in the region are Pellaea glabella, Poly- podium virginianum, Asplenium trichomanes, and Camptosorus rhizophyllus. Boreal species associated with bogs and cool swamps are Equisetum pratense, Equisetum scirpoides, and Equisetum sylvaticum; Lycopodium — an- notinum, Lycopodium clavatum, Lycopodium inundatum, Lycopodium lucidulum, and Lyco- podium obscurum; also, Phegopteris connectilis (after Thaler and Plowright 1973). A species of northern Appalachian affinity is Dryopteris goldiana. Woodwardia virginica is a fern with Atlantic coastal plain affinities (Cody 1963). These local pteridophytes which may be con- sidered essentially southern are Equisetum laevigatum, Selaginella apoda, (Phegopteris hexagonoptera — a species excluded from the Waterloo list), Athyrium pycnocarpon, and Asplenium platyneuron. Thus in its pteridophyte flora as in its other floristic origins, the Waterloo region exhibits an overlap between northern and southern plants. Acknowledgments In addition to the many people whose names appear throughout the paper, we thank the curators of various herbaria for allowing us to examine specimens. R. L. Hauke, University of Rhode Island and W. H. Wagner Jr., University of Michigan, kindly determined many of our Equisetum and Botrychium specimens, respec- nels AXy WW awlor, ID), Jb. Ibeslie, aimel Jj. Ledingham of Cambridge (Galt) helped greatly THE CANADIAN FIELD-NATURALIST Vol. 91 in locating Herriott’s collecting sites. R.A. MacLaren, of Hamilton, V.C. Shantz of Kitchener, and D. Brown of University of Waterloo contributed records of several rare species; P. F. J. Eagles of University of Water- loo advised us on some locations. The field assistance to Campbell by A. A. Reznicek (Erindale College), G. R. Donaldson (Ecoplans Ltd., Waterloo), and J. L. Campbell is gratefully acknowledged. D. L. Campbell assisted with field work and compilation of records, and typed the manuscript. Literature Cited Cody, W. J. 1963. Woodwardia in Canada. American Fern Journal 53: 17-27. Crabbe, J. A., A. C. Jermy, and J. T. Mickel. 1975. A new generic sequence for the pteridophyte herbarium. Fern Gazette 11(2, 3): 141-162. Cruise, James E. 1969. A floristic study of Norfolk County, Ontario. Transactions of the Royal Canadian Institute 35: 3-116. Fernald, M. L. 1950. Gray’s manual of botany. 8th edition. American Book Company, New York. Gleason, H.S. 1952. The new Britton and Brown illus- trated flora of the northeastern United States and adjacent Canada. Hafner Publishing Company, New York. Montgomery, F. H. 1944. A botanical survey of Waterloo County, Ontario. M.A. thesis, McMaster University, Hamilton. Montgomery, F. H. 1945. A botanical survey of Waterloo County, Ontario. Transactions of the Royal Canadian Institute 25(2): 217-266. Taylor, T. M. C. 1934. Notes on some Ontario pterido- phytes. American Fern Journal 24(3): 79-83. Thaler, G. R. and R. C. Plowright. 1973. An examination of the floristic zone concept with special reference to the northern limit of the Carolinian zone in southern Ontario. Canadian Journal of Botany 51(10): 1765-1789. Wherry, E. T. 1961. The fern guide: northeastern and mid- land United States and adjacent Canada. Doubleday and Company, Garden City, New York. Widén, C.-J. and D. M. Britton. 1971. Chemotaxonomic investigations on the Dryopteris cristata complex in North America. Canadian Journal of Botany 49(7): 1141-1154. Received 4 February 1977 Accepted 18 June 1977 The Biological Flora of Canada — A New Series GEORGE H. LA ROI Department of Botany, University of Alberta, Edmonton, Alberta T6G 2E9 La Roi, George H. 1977. The biological flora of Canada — A New series. Canadian Field-Naturalist 91(3): 269-272. Abstract. The Canadian Field-Naturalist will publish a continuing series of papers on the biology and ecological life history of vascular plant species native or well naturalized in the flora of Canada. This paper sets forth the guidelines and format for contributing authors. The new Canadian series is patterned after the Biological Flora of the British Isles, a continuing series published in the Journal of Ecology. Notice to Contributors The Canadian Field- Naturalist has agreed to publish a continuing series of papers on the biology and ecological life history of vascular plant species that are native or well naturalized in the flora of Canada. These accounts will be written by various authors, follow approved guidelines and format (see below), conform to the standards and style of this journal, and include distribution maps, literature citations, and other information needed to describe and interpret the role of the species in the vegetation of northern North America. This is usually best achieved by a synthesis of information obtained from the literature together with original research on aspects not previously investigated. The accounts will be numbered and appear in order of acceptance, with a maximum of two published per issue. A similar project, The Biological Flora of the British Isles, was undertaken by the British Ecological Society in 1945 and has been a regular feature of The Journal of Ecology ever since (British Ecological Society 1975). Recently the Canadian Journal of Plant Science has initiated a series on The Biology of Canadian Weeds (Cavers and Mulligan 1972) and one on The Biological Flora of the Canadian Prairie Provinces (Looman 1973). Species already registered, or previously published, or more appropriately published in the latter two series will not be registered for publication in our series; close liaison is maintained between editors of the two journals to insure that the same species is not registered in more than one series. Inquiries about the series and offers of contributions should be made to the author; the latter are subject to approval by a committee of botanical associate editors. Finished manuscripts may not exceed 10 journal pages, including text, tables, and illustrations; they should be submitted to the Editor. Guidelines for Contributors Only papers on vascular plants will be accepted, including aquatic, wetland, and terrestrial species; this restriction will be reviewed periodically. Very closely related species may be treated together in some cases. Both native and very well naturalized species are eligible. Species already treated in the Biological Flora of the British Isles (British Ecological Society 1975), Silvics of Forest Trees of the United States (Fowells 1965), and equivalent references will be registered, written and published as “Supplementary Accounts” to avoid needless replication. Decisions on acceptance of a particular species will be influenced by (a) its ecological and economic importance in Canada, (b) extent of its total range in Canada, e.g., species reaching their northern limits in the extreme south of Canada will receive lower priority. Acceptance of offers and acceptance of manuscripts are entirely separate matters. Poorly known species should not be offered unless the contributor is able and willing to obtain the substantive information specified in the schedule below. Contributors will be allowed to register only two species in our list of species in preparation, as single or multiple authors. Approved offers will be registered for three years, after which reapproval must be sought or the species will be deleted from the list and made available to other potential 269 DUG) THE CANADIAN FIELD-NATURALIST Vol. 91 contributors. An approved blank base map will be provided to approved contributors; it will cover all of Alaska, Canada, Greenland, St. Pierre and Miquelon, and the northern fringe of the conterminous United States. Contributors must consult with and acknowledge the Herbaria of the Canada Department of Agriculture and the National Museum of Canada in Ottawa regarding loans of specimens and use of their facilities for checking determinations of specimens and plotting distributions of species on dot maps. Subsequent papers reporting significant range corrections and extensions, and submitted to this journal, should employ the same map and clearly depict the changes. Schedule for Contributors (1977) 1. Name.' The scientific name (genus, species, authority) currently accepted, followed by the subgenus or section, and family. No more than three of the most important synonyms. English and French vernacular names most commonly used in Canada. 2. Description of the Mature Plant. Concise description of the adult sporophyte, with attention to (a) Raunkiaer life-form and perennation, (b) Shoot morphology, i.e., above-ground vegetative structures, (c) Root morphology, i.e., below-ground structures, (d) Inflorescence. Include diagnostic and important characters. Important infraspecific morphological variation within the region including (e) Subspecies, (f) Varieties (g) Ecotypes, (h) Chromosome number(s) of species together with locality and authority. Photographs and well-executed drawing of typical living and mounted herbarium specimens. 3. Distribution and Abundance. Native or introduced since European settlement, with notes on first records, rate of migration, and stability of present distribution. (a) Geographic range of species and its major variants in region from herbarium records confirmed by the author(s) and plotted on a standard base map to be provided; notes on abundance in region; maps and notes on distribution outside the region in less detail; cite published distribution maps. (6) A/titudinal range in and outside region; notes on abundance and infraspecific variation in relation to elevation. 4. Physical Habitat. (a) Climatic relations in general, then specific ranges and optima for temperature, precipitation, humidity, light, wind, etc., with attention to seasonal variations of these in relation to the life cycle and phenology of the plant. (6) Physiographic relations, including slope and exposure, ranges of geologic parent materials, soil types, drainage classes and moisture regimes; soil terminology should conform to the current edition of The System of Soil Classification for Canada (Canada Department of Agriculture 1974). (c) Nutrient and water relations, including general requirements, performance in specific nutrient and moisture regimes, and the plant’s role in the total nutrient and water balances of the ecosystem. Emphasize specific factors that exert primary control on distribution and abundance in the region. 5. Plant Communities. General summary of plant community types in which the species occurs in the region, with notes on local distribution, abundance, and reproductive success in them. A species- stand table showing the physiognomy and floristic composition of a representative spectrum of community types in which the species is an important component and/or grows well. Cite publications containing community descriptions and tabulations which include the species. 6. Growth and Development. Concise description of plant growth and development from germination through maturity to senescence and death. (a) Morphology of plant at critical developmental and phenological stages depicted in well-executed drawings, with notes on morphological adaptations, rates of growth, and duration of stages. (b) Physiology of plant at critical developmental and phenological stages depicted in tables giving the rates of vital processes (photosynthesis, respiration, translocation, transpiration) and normal physiological states (leaf water potential, biochemical composition) under specified conditions of light, moisture, nutrients, 'Each account will be given a serial number when accepted for publication. 1977 LA ROI. BIOLOGICAL FLORA OF CANADA Dal and temperature. (c) Phenology of species at different localities within region including dates of germination; onset of vegetative growth, flowering; maturation and dispersal of seeds; peak root and shoot growth; return to dormant state above and below ground. 7. Reproduction. (a) Floral biology of the plant including mode of pollination; incidence of autogamy, allogamy, agamospermy, vivipary. (b) Seed production and dispersal, including data on age-specific numbers of seed per fruit and per plant; modes of dispersal and their effectiveness; seed production in relation to geographic location, habitat quality, year. (c) Seed viability and germination under different conditions, with notes on the nature of dormancy and conditions necessary for maintaining or breaking it; specify methods of seed collection, storage, and treatment as well as germination conditions. (d) Vegetative reproduction mode(s) and rates in relation to geographic location, habitat quality, age of plant; importance compared to sexual reproduction. 8. Population Structure and Dynamics. (a) Dispersion patterns at different stages in the life cycle and in different habitats. (6) Age distribution, age-specific mortality rates, and longevity in different habitats, with notes on causes of mortality. (c) Size distribution of individuals in populations of different habitats, with data on correlations between age and size of individuals. (d) Growth and turnover rates of stable and unstable populations in different habitats, expressed as percentage increase, decrease and replacement of population (density, biomass, etc.) per year. (e) Successional role of species in relation to geographic location, habitat quality, type of disturbance, severity of disturbance. 9. Interaction with Other Species. Concise description of (a) Competition with com- monly associated plant species for light, nutrients, water and space; (b) Symbiosis with other organisms; list important pollinators, endophytic nitrogen-fixing bacteria, mycorrhizal species; (c) Predation and parasitism with other organisms; list host species if the plant is parasitic; list important animal and fungal predators and parasites, and bacteria and virus disease organisms for which the plant is prey or host, the parts and/ or life cycle stages used by them, their effects and symptoms, and their geographic distribution and abundance in relation to those of the plant; (d) Toxicity and allelopathy to other organisms; list substances and their effects on other species. 10. Evolution and Migration. A brief account of the origin and phylogeny of the species, and its history as a member of the Canadian flora. Notes on fossil records, glacial refugia, migrational patterns, etc. If introduced since European settlement, provide relevant details in (3). Chemotaxonomic, cytotaxonomic and other supporting evidence may be cited here. Occurrence and frequency of hybrids with other species, how they may be recognized, and where they may be found. 11. Response Behavior. Normal responses of individuals and populations to (a) Fire, (b) Grazing and harvesting, (c) Flooding, (d) Drought, (e) Herbicides, (f) Chemical changes including toxic compounds and nutrient levels, (g) Other factors. Significant adaptations should be noted. 12. Relationship to Man. Concise summary of the plant’s importance to man, as a crop, pest, or reclamation species, etc., in the past and present in the region. Man’s influence on the distribution and abundance of the species in the region. 13. Special Features. Noteworthy information not provided for elsewhere in the schedule. Acknowledgments The author thanks G. W. Argus, C. D. Bird, W. J. Cody, and L. C. Smith for their suggestions and constructive criticisms of the guidelines and schedule. Some Relevant References Arno, S.F. and J.R. Habeck. 1972. Ecology of alpine larch (Larix J/yallii Parl.) in the Pacific Northwest. Ecological Monographs 42: 417-450. Bliss, L. C. 1971. Arctic and alpine plant life cycles. Annual Review of Ecology and Systematics 2: 405-438. DUP THE CANADIAN FIELD-NATURALIST Vol. 91 British Ecological Society. 1975. The biological flora of the British Isles: Notice and revised schedule for contributors. Journal of Ecology 63: 335-344. Canada Department of Agriculture. 1974. The system of soil classification for Canada. Queen’s Printer, Ottawa. Cavers, P. B. and G. A. Mulligan. 1972. A new series — The biology of Canadian weeds. Canadian Journal of Plant Science 52: 651-654. Fowells, E.H. 1965. Silvics of forest trees of the United States. United States Department of Agriculture, Agriculture Handbook Number 271. 762 pp. Hall, I. V., L. P. Jackson, and C. F. Everett. 1973. The biology of Canadian weeds. 1. Ka/mia angustifolia L. Canadian Journal of Plant Science 53: 865-873. Looman, J. 1973. Biological flora of the Canadian Prairie Provinces. I. Oxytropis besseyi (Rydb.) Blank. Canadian Journal of Plant Science 53: 677-687. Matthews, V. B. and P. W. Conrad. 1968. An ecological life history of tall bluebell (Mertensia arizonica var. leonardi) in Utah. Ecology 49: 1113-1123. Morgan, M.D. 1971. Life history and energy relationships of Hydrophyllum appendiculatum. Ecological Monographs 41: 329-349. Pelton, J. 1951. Outline for ecological life history studies in trees, shrubs, and stem succulents. Ecology 32: 334-343. Pelton, J. 1953. Ecological life cycle of seed plants. Ecology 34: 619-628. Penfound, W. T. 1952. An outline for ecological life history studies of herbaceous vascular hydrophytes. Ecology 33: 123-128. Richards, P. W. 1945. The biological flora of the British Isles. Chronica Botanica 9: 140-144. Rudolf, P.O. 1958. Silvical characteristics of jack pine (Pinus banksiana Lamb).United States Department of Agriculture, Forest Service, Lake States Forest Experiment Station Paper Number 61. 31 pp. Stevens, O. A. and L. Rock. 1952. Outline for ecological life history studies of herbaceous plants. Ecology 33: 415-422. Wein, R. W. 1973. Eriophorum vaginatum L. Biological flora of the British Isles. Journal of Ecology 61: 610-615. Received 24 June 1977 Accepted 10 July 1977 Movements and Habitat Use among Interacting Peromyscus leucopus as Revealed by Radiotelemetry DALE M. MADISON Department of Biology, State University of New York at Binghamton, Binghamton, New York 13901 Madison, Dale M. 1977. Movements and habitat use among interacting Peromyscus leucopus as revealed by radiotelemetry. Canadian Field-Naturalist 91(3): 273-281. Abstract. Radiotelemetry and trapping methods were used in a systematic effort to record habitat use, home range size, and social interactions in a free-ranging population of Peromyscus leucopus. A total of 64 mice were censused in an isolated 2.3-ha woodlot in May and June 1975, revealing a large spring density of 21.7 resident mice/ha. The sex ratio in each age class was approximately I:1. The subadult age class had significantly more transient individuals than did the juvenile or adult age classes. From the 36 adults captured in traps, nine males and six females were followed using radiotelemetry during an 8-day period. These mice were located once day and night for a total of 124 detections. Daytime refuges were usually in groundhog (Marmota monax) burrows and rock piles, and were similar for both sexes. Nocturnal activity was at ground level, with the males showing a significant tendency to occupy the surrounding fields instead of the woodlands. The occupation of defecation chambers in the groundhog burrows and the caching of large fecal pellets in other refuges suggested interspecific coprophagy. The large number of hickory nuts and nut fragments in the refuges suggested abundant food supplies during the winter. A large winter food-supply together with the abundant rock piles and groundhog burrows probably accounts for the large spring density in the population studied. Home range size (minimum area method) was similar for both sexes, averaging 0.1 ha. Sustained social contact was frequent between adult male and female voles, whereas low levels of contact occurred between adults of the same sex, and then only between males. These data support the potential of radiotelemetry to supply important biological information on the movements of free-ranging rodents. Key words: Peromyscus, habitat, movements, radiotelemetry, spacing, nests Rodent studies have been of central impor- tance to the formulation of many ecological principles. Yet little direct information is available on daily activities and resource use in free-ranging rodent populations, even in such a widely studied species as Peromyscus leucopus (Metzgar 1971: Myton 1974; Terman 1968). Historically, methods involving trapping, pho- tographic, and smoked-paper techniques have limited attention to positions in space or time selected by the experimenter, not by the animal. More recent rodent studies involving radio- telemetry (Banks et al. 1975; Brooks and Banks 1971: Chute et al. 1974; Madison 1977; Mineau and Madison 1977: Shields 1976) or radio- isotope tracking (Ambrose 1969, 1973: Graham 1968) are beginning to yield direct information on natural patterns of movement. A study such as that of Banks et al. (1975) involving the simultaneous tracking of a large number of free- ranging individuals is important since it provides accurate information on both individual and group (population) responses to environmental and social variables on a daily basis. The following study applies simultaneous radio- tracking procedures in a study of the move- ments and habitat use of free-ranging P. leucopus, a small, highly mobile rodent species. Methods Study Area The study took place in an isolated 2.3-ha (5.3- acre) deciduous woodland near Front Royal, Virginia in 1975. The surrounding land was either alfalfa, hay, or early seral, secondary succession fields. The nearest point to another section of woodland was approximately 250 m north and east of the study woods. The woodlot had a dense stand of hickory (Carya sp.) and a large variety of oaks (Quercus spp.). The area was browsed by deer as could be noticed by the reduced vegetation near ground level. On several occasions, deer were actually observed in the study area at night. The area was also extensively used by groundhogs (Marmota monax); 13 burrow systems with a total of 36 entrances were counted within the study area. Most of these systems were active. Six rockpiles were dis- tributed toward the periphery of the study area, ranging in size from 0.5m highX2m in diameter to 1.5 m high X 3 m wide X 10 m long. These were probably created when the adjacent fields were cleared for cultivation. Grass and succulent broadleaf vegetation invaded the woods 30m on the northwestern edge of the woodland, but otherwise stopped abruptly at the woods’ edge. There was a fairly tegular incline of Jnif3) 274 20 degrees from southeast to northwest through the area. Trapping Techniques On 21 May, trapping was initiated using 21 Longworth live traps, both to census the population and to provide individuals for radio- tracking. Three trap lines were established about 40 m apart, with the distance between traps in each line averaging 25m. The traps were supplied with rolled oats for bait and a paper towel for bedding. The traps were set before dark, checked around 2200 hours, reset, and then checked around 0100 hours. This pro- cedure continued until essentially all the adult mice were marked (4 June). Then radiotracking began and only the 2200-hours trap check was made every other day until 20 June when the study was terminated. Six additional traps were added after 10 June to facilitate the recapture of mice with transmitters. Upon initial capture, each animal was weighed, given a unique toe clip combination, checked for sex and reproductive condition, and then released. During subsequent captures, the procedure was repeated except for toe-clipping. Dramatic weight loss in the females was used as an index of parturition (four females in this study lost from 7 to 10 g between successive trap checks). Radiotelemetry Techniques All the adult mice (20 g or more) captured from 2 to 4 June were given transmitters and then tracked concurrently. Conventional radio- telemetry equipment was used (AVM Instru- ment Co., Champaign, Illinois), including SM-1 radiotransmitters with internal antennae, mul- tiple channel LA-12 radioreceivers, and hand- held 4-element Yagi antennae. Details on the radiotelemetry equipment and collar attachment are available in Mineau and Madison (1977). The positions of the mice were determined by a single observer. During the day the mice occupied nests and therefore could be ap- proached directly to obtain a position. At night, a 6-volt head lamp and a more cautious method of approach were used. Typically, the area to be searched was scanned with the receiving antenna from about 30-40 m distance to assess the different mice present. Then the nearest mouse THE CANADIAN FIELD-NATURALIST Vol. 91 was approached, first tangentially to obtain a rough “triangular” fix, then more directly to within about 10 m. At this distance, the direction of maximum signal intensity was scanned slowly by light. Often the light revealed the animal either moving on the woodland litter or perched on some surface object. A coded marker was attached nearby, and the position was described in a notebook in full. In cases where two signals were in close proximity, the observer constantly switched between the frequencies during the approach to assess carefully the proximity of the animals to each other. There was little evidence that the mice were fleeing from the observer. Rather, immobility or slow “foraging” move- ments were typical. Radiotelemetry positions were recorded at least once both day and night, thus obtaining one measure of nest location and usually one of surface activity each 24-h period. Transmitter signals at night from day nest positions usually indicated the loss of transmitter collars, except in the case of females tending litters. If a transmitter was no longer on an animal, the transmitter (usually in a nest) was retrieved and the nest was examined. During the study, many transmitters stopped functioning. The problem was that conspecifics gnawed through the epoxy coating of the transmitter, often destroying the transmitters. That mutual grooming, and not self-grooming, was responsible for transmitter and collar damage was verified in cage studies where only those mice housed together showed such damage. These problems could have been avoided, if anticipated, by using dental acrylic instead of epoxy, and by substituting a different collar material for the cloth used. This damage was not encountered in extensive studies on Microtus pennsylvanicus (Madison, unpub- lished) and on a low-density population of P. leucopus (Mineau and Madison 1977). Accord- ing to the importance of mutual grooming in mating and in the establishment and main- tenance of dominance relationships in Pero- myscus (Eisenberg 1968), transmitter-collar damage is not very surprising. Because sustained proximity and gnawing is required to damage the transmitters, and because overt aggressive actions would presumably result in one mouse LMT. fleeing from the other, social interaction was considered to have been measured. Data Analysis All the information from trapping and radiotelemetry will be used during the analysis, but two clarifications are necessary. First, unless stated otherwise, only the mice captured and determined to be “residents” (see “Population structure”) will be considered in the analysis. Second, although 19 mice were given trans- mitters, four were never located using radio- telemetry. Three of these were eventually recaptured without transmitter collars. Of the 15 mice actually monitored, two were tracked only for the first day before their transmitters were damaged. The remaining 13 were tracked both day and night. Thus, the radiotelemetry analyses include either 13 or 15 individuals, as is relevant to the specific analysis. Population structure. In the population studied, the age class criteria of Bendell (1959) and Snyder (1956) were used. Residency was assumed for a mouse if it was captured 2 or more times over at least a 4-day period. All other mice were considered transients. The 4-day criterion was necessary to distinguish between a resident, who is captured only a few times over a long time interval because of a small or peripheral home range, and atransient, who may also be captured a few times, but only in close succession while passing through the study area. The relative numbers of transient and resident mice in the different age classes were compared using the Fisher exact test (Sokal and Rohlf 1969). This test gives exact probabilities and 1s particularly useful for small sample sizes and small expected frequencies under the null hypothesis. Resource use. Data on shelter and food use were collected from the observations on the adult mice that were monitored using radio- telemetry. The day shelters or refuges were classified as either a small ground burrow (mouse-sized or chipmunk-sized entrance), a groundhog burrow, a rock pile, or a tree. Differences in shelter or refuge use between the sexes for the day and night positions were analyzed using Fisher’s exact test (Sokal and Rohlf 1969). The utilization of certain foods was estimated from the contents of day nests. MADISON: PEROMYSCUS MOVEMENTS AND HABITAT USE BY TELEMETRY 275 Home range size. Home range size was determined only for the adult mice that were radio-tracked. For the estimates, the trap locations and radiotelemetry positions were enclosed within a peripheral line (minimum area method). Each trap location is arbitrarily considered to be a circle with a 3-m radius, instead of a radius one-half the distance to the nearest trap as in the boundary strip method. This 3-m distance was chosen because of the variable distance between traps and because the smallest interval between the 21 traps set was 6m. No attempt was made to adjust for “unoccupied regions” within the peripheral lines, because the period of study was short and the observed home ranges were relatively compact. Social interactions. Estimates of social inter- action and tolerance were obtained by con- sidering the instances of transmitter-collar damage, double capture, and social encounter and nest cohabitation of mice tracked with radiotelemetry. The double captures in traps represented close following of one mouse by another, because the trip mechanism in each trap had been adjusted for extreme sensitivity. In cases of social intolerance (e.g., one mouse chasing another) it is assumed that the fleeing mouse would not enter a trap as a means of escape. In no case was there any evidence of fighting by the mice captured together. Results A total of 63 P. leucopus (49 residents) were captured in traps in the study area. The residents were captured on 290 occasions (average of 5.8 captures/mouse). The 15 mice tracked were trapped on 124 occasions, and from the beginning of radiotracking had 64 day and 44 night positions recorded (Table 1). The animals were tracked for an average duration of 5 days each (1 to 8 day range). On only three occasions did we fail to find a mouse whose transmitter, upon subsequent inspection, was known to be functioning. On two of these occasions, which were recorded successively for the same animal, the failure probably resulted from a weak transmitter signal combined with a shift in habitation to a groundhog burrow system. 276 THE CANADIAN FIELD-NATURALIST Vol. 91 TABLE 1—Capture, position, and home range data for Peromyscus leucopus tracked with radiotelemetry Number Number Total Number Home trap Number radio number days range captures traps positions positions tracked (acres)! Males (N = 9) Mean Ws DP) 7.6 15.2 48 0.24 SD 3.4 0.7 4.0 2.9 2.1 0.14 Range 3-13 1-3 1-14 12-20 1-8 0.04-0.43 Females (N = 6) Mean 9.2 2.8 6.7 15.8 5.0 0.26 SD 32 1.8 3.4 7.3 D3 0.24 Range 2-15 1-5, 1-11 3-24 1-7 0.08—0.64 'The home range calculations for the females are based on N = 5; one female was omitted from the calculations because of inadequate data. Population Structure The density of the resident population was 21.3 individuals/ha (8.4 mice/acre). Three different age class modes were observed, one averaging 13 g representing juveniles about 4 weeks of age, a second averaging 16 g repre- senting subadults about 7 weeks of age, and a third around 23 g representing sexually mature adults of a wide age spectrum (Figure |). The sex ratio for the resident immature mice was approximately I:1 (8 males, 9 females). The ratio for the adult residents was similar (17 males, 15 females). Transient status was recorded for 2 of 13 juveniles, 8 of 14 subadults, and 4 of 36 adults. The two mice at 14 g were divided among the ) Number of animals (6) juvenile and subadult age groups because of unclear group affiliation. The hypothesis of no difference in the relative numbers of resident and transient mice among the juvenile and subadult age groups is rejected (P= 0.03), as is the null hypothesis relative to the subadult and adult age groups (P< 0.001). The null hypothesis relative to the juvenile and adult age groups, however, is not rejected. These results indicate more transient mice, and hence a greater degree of either dispersal or death, in the subadult age class. Habitat Use Refuge use (rock piles, groundhog burrows) at RIRIEIEI oud oud ou) Ba) Ba) |= [) Nee] 30 eras echt (gna ae FiGure |. Weight frequency distribution of all Peromyscus leucopus captured in the study area. Circled F (female) and M (male) indicate transient individuals. Solid dots indicate pregnant females; open dots, females having had a litter within the study period. 1977 MADISON: PEROMYSCUS MOVEMENTS AND HABITAT USE BY TELEMETRY AT) TABLE 2—Habitat use during night positions (2100-0400 hours) for 13 Peromyscus leucopus tracked using radiotelemetry in June 1975. Number of different occurrences are given. Numbers of different animals appear in parentheses Tree aime neal (a) Leaf Groundhog Rock Sex 0-1.4 1.5-2.9 3.0+ Grass litter burrow pile Males 27(8) 2(2) 2(2) 0(0) 13(5) 8(6) 0(0) 2(1) Females 17(5) 5(4) I(1) I(1) 0(0) 5(4) 3(2) 2(1) Totals 11(7) 13(5) 13(10) 3(2) 4(2) night occurred on 5 of 17 occasions for females, but only on 2 of 27 occasions for males (P = 0.06; Table 2). Nocturnal activity was primarily at ground level (leaf litter, grass, tree bases) for both sexes. During ground level activity, however, the males occupied the grass areas of the surrounding fields on 13 of 23 occasions compared to 0 of 10 occasions for the females. The hypothesis of no relation between the use of the grass fields and the sex of the adult 1s rejected (P=0.002). These results suggest differences between the sexes in habitat preference and (or) food utilization at night. Refuge use by the 15 mice tracked during the day consisted primarily of groundhog burrows, with 10 of 15 mice found in these burrows on 34 of 64 occasions (Table 3). Rock piles were preferred next with six mice in these structures on [5 occasions. About 50% (8 of 15) of the mice utilized more than one day refuge during the study. There was no indication of differences in day refuge preference between the sexes. The utilization of certain foods was suggested from the contents of the six day nests that were excavated to retrieve transmitter collars. Three units were found in the defecation chambers of groundhog burrows. A prominent feature of these chambers was a layering of groundhog feces and leaves with a thickness of 0.2-0.3 min the chamber floor, giving a compost effect with the associated increase in temperature (the warmer temperature was gauged by touching). The topmost fecal pellets were green as if consisting of semi-digested grass mulch. The possibility of these pellets serving as a supple- mentary food source is suggested (interspecific coprophagy). In two of these groundhog refuges, a large cache of nuts, seeds, and shell fragments (e.g., hickory, cherry) were uncovered, but no nest structure was found. Two other transmitters were recovered from a single refuge site ina rock pile. This refuge contained a fur-lined grass nest, a large cache of nut and nut fragments (hickory, cherry, white oak acorns), gnawed twigs (cherry (Prunus sp.), birch (Betula sp.)), male flower parts from oaks (Quercus spp.), legumes, several Coleoptera insect parts, and one area with a concentration of mice feces. Notable among the nest site contents were several bone fragments scarred by Peromyscus tooth marks, and two fecal pellets (one a 2-cm-wide X I-cm-long segment from a large herbivorous mammal, and one a 0.7-cm-wide X 3-cm-long scat containing hair and nut fragments from a small carnivore). The sixth transmitter was found ina cavity at the base of a dead cherry tree. A small cache of leaf and nut parts was found, witha nut composition and tooth marks similar to that in the rock pile nest. TasL_eE 3—Refuge use during day positions (0700-1900 hours) for 15 Peromyscus leucopus tracked using radio- telemetry in June 1975. Number of different occurrences are given. Numbers of different animals appear in parentheses Ground- Small hog ground Rock Sex burrow burrow pile Tree Males 40(9) 23(6) 2(1) 8(4) 7(2) Females 24(6) 11(4) 4(3) 7(2) 2(2) Totals 34(10) 6(4) 15(6) 9(4) THE CANADIAN FIELD-NATURALIST Vol. 91 ——— metres FiGuRE 2. Home ranges of all resident adult P. /ewcopus that were radio-tracked. The radiotelemetry positions (small circles; solid = night, open = day) and trap capture positions (large circles) are indicated for the males (A) and females (B). The letters next to the radiotelemetry positions identify refuges in trees (T), groundhog burrows (G), small ground burrows (S), and rock piles (R). The nursing females are indicated by N. The small arrows associated with radio- telemetry positions in areas of home range overlap point to the home range to which the positions belong. Home Range Size Male home ranges averaged 0.10 ha (0.24 acre, SD = 0.14 acre) based on 12 to 20 capture and radiotelemetry positions for each animal (Figure 2, Table 1). For the females, an average area of 0.1] ha (0.26 acre, SD=0.24) was recorded based on 14 to 24 positions per animal. The small difference in home area size between the males and females was not significant. These estimates are independent of “edge” effects since the animals living in the border areas were tracked into adjacent fields. Social Interaction There were 15 instances where the fate of the transmitter and collar was known, with 13 units sustaining damage (86.7%) within 8 days (mode of 4) of the initial collar attachment. Only two mice retained their collars and transmitters throughout the tracking period without damage. These data indicate frequent social interactions of a sustained nature within the population. The eight instances of double capture also support the occurrence of social interactions within the population (Table 4). Four double captures involved only juveniles. Because of the similarity in weights within each pair, it is suggested that litter mates were travelling together. Two scrotal males were involved in the remaining four double captures. These males NOW TABLE 4—Instances of double captures of Peromyscus leucopus in traps. The weights are those recorded at time of double capture Animals captured together Date Sex Weight(g) Sex Weight (g) 1 June M, 22 F 25 2 June same Ms 23 3 June M, 23 In 20 10 June same M 16 26 May M 15 M 13 10 June F 9 M 10 17 June F 13 If 15 20 June F 14 F 15 s = scrotal; ns = nonscrotal; | = litter had lost their transmitter collars on the same night as transmitter attachment, thus suggesting more extensive involvement in social inter- actions for these males. The radiotelemetry data reveal nine instances of adult mice occupying similar locations at the same or different times (Table 5). Eight of the TABLE 5—Instances of concurrent or sequential occupation of a given location by Peromyscus leucopus bearing radio- transmitters. The mice listed in the first column previously occupied the location, unless both mice were observed at the location for the first time (indicated by ). GHB = groundhog burrow, , denotes a nursing female; each number identifies an individual Animals Num- Num- Date per SO ele SOX Particulars 4 June 1 M? 2 F Concurrent, rock pile Concurrent, GHB, within 2m Sequential, tree nest Concurrent, GHB Concurrent, GHB Sequential, GHB Sequential, GHB Concurrent, leaf litter, within 6m Sequential, GHB 5 June 3 M 5 M 5 June 3 F, 4 6June 4 \sh 4 7June 4 JB 4 8 June 4 F, 4 8 June 4 F, 4 Se Se Sa 8 June 3 JR 4 9June 4 En 4 K MADISON: PEROMYSCUS MOVEMENTS AND HABITAT USE BY TELEMETRY 279 nine instances involved mice of the opposite sex; the remaining instance involved two males 2 m apart in the same groundhog burrow system. In total, not one instance of double capture or cohabitation between adults involved two females. Three such instances occurred between males, while 10 instances occurred between mice of opposite sex. Discussion Several observations on the numbers, move- ments, and habitat use of P. /ewcopus in this study were unexpected, although the pre- liminary information on the intensity and specificity of social interactions agreed with expectations. The first unexpected finding was the large spring density of resident P. /eucopus of 8.4 mice/acre. This compares to normal values of 4 mice/acre (see Terman 1968; Jackson 1961). Although not excessively above average values, the observed density was conspicuous since the habitat structure (1.e., the scarcity of low-level vegetation and hence reduced ground-level cover and food resources) would normally lead to predictions of low density (e.g., see Bendel 1959; Myton 1974; Stickel and Warbach 1960). Two features of the habitat are believed to have been responsible for the high spring density. First, the dense stand of hickory trees and the conspicuous scattering of nuts on the woodland litter may have supplied a large fall harvest of nuts for winter food needs. The number of shell fragments of hickory nuts, scarred in the characteristic way for Pero- myscus (Eisenberg 1968; Nicholson 1941), were by far the most common item in the excavated nests. Second, the abundance of groundhog burrow systems, both abandoned and occupied, probably supplied a considerable number of excellent subterranean retreats for winter and spring shelter. Subterranean nests are favored by P. leucopus during the winter (Nicholson 1941). It is not known to what degree the closure of burrows by groundhogs during hibernation affects burrow availability during mid-winter. Co-residence by Marmota monax and P. leucopus in Marmota burrow systems, in- cluding the accumulation of food items by P. leucopus, is the first report of such burrow occupancy. The possibility of reproductive, thermoregulatory, and energetic gains 1s clear, as 280 can be deduced from the studies of Millar (1975) and Dudley (1974). Another unexpected finding was the degree of ground-level activity and the frequent use of subterranean and rock-pile refuges. The habitat, as previously mentioned, was conspicuously open, being influenced primarily by deer grazing. Fallen logs and branches were evident, although not sufficient to obscure vision at ground or eye level for at least 70 m. Previous reports suggested that nesting and foraging in trees should have been more frequent, especially during the spring and summer (Horner 1954; M’Closkey 1975, 1976; M’Closkey and Lajoie 1975; Nicholson 1941). That the transmitters did not cause the ground-level activity is supported by the observations of tree-nest use by P. leucopus bearing transmitters (Mineau and Madison 1977). In addition, when the mice were observed in trees, or even when seen running over surface rocks and fallen logs and branches, they did not appear to have problems with balance, agility, or with gripping the substrate or tree bark. Finally, the groundhog, rock-pile, and other ground-level nests showed evidence of long-term occupancy (old nut fragments, layer- ing of old nest material) beginning long before the period of this study. A third unexpected finding was the tendency of male P. leucopus to move into the surround- ing fields during much of their nightly (for- aging?) activity, but to show the same preference as the females in the type of daytime retreat. The possibility exists that by chance more males than females were selected from areas near fields for radiotracking. The use of grass or early sec- ondary succession fields by P. leucopus for foraging has been reported (e.g., Stickel 1968; Whitaker 1966; Pearson 1959), but the relative frequency of use of these fields by mice of different sex has not been examined. The flexibility in diet of P. Jewcopus (Whitaker 1966; Drickamer 1972, 1976) may allow temporary partitioning of the food supply when the spatial intolerance of one sex exceeds that of the other. Hence, nursing or pregnant females could be particularly intolerant of conspecifics and could partially exclude males and non-reproductive females to less favorable areas in the habitat. In the case of isolated woodlots, these less THE CANADIAN FIELD-NATURALIST Vol. 91 favorable areas could be the surrounding fields. The possibility of sex differences in diet or habitat use certainly needs additional study before further comments are warranted. Given the presence of a higher density popu- lation of P. leucopus, the findings relative to social interactions agree with previous studies. First, the higher frequency of transient mice among the subadult age class is consistent with earlier reports (see Terman 1968), and suggests that not much space exists in the resident population for young reproductives. Second, despite the higher density, the home range sizes are in the range of previous estimates (Terman 1968), and this agreement supports size stability of home range size at different population den- sities (Metzgar 1971). The much larger 3.1- and 2.3-acre home ranges recorded for transmittered P. leucopus ina low-density population (Mineau and Madison 1977) suggests that home range size may vary with extremes in population density, and that the transmitters did not depress home range size in the present study. Third, the preliminary indication of sustained social inter- action between mice of the opposite sex, in comparison to mice of the same sex, affirms the observations of Metzgar (1971). Thus, in conclusion, the potential of radio- telemetry for obtaining more precise informa- tion on many aspects of the biology of free- ranging rodents is clear. Even the relatively short term of this study was sufficient to provide new and potentially significant information for P. leucopus concerning interspecific coprophagy, interspecific nest cohabitation, and temporary or partial habitat (food?) partitioning between adult males and females. Acknowledgments I am especially grateful to Chris Wemmer for making facilities available for the study at the Center of Conservation and Research, Front Royal, Virginia. I thank Bruce Webster, Melissa Ditton, Geoffrey Gartshore, and Heather Ham- ilton for their help in the field. Peter Grant, Donald Kramer, and Wolfgang Schleidt made helpful criticisms during manuscript prepara- tion. This study was supported by NRCC grant A-9591 and by McGill University. MIT Literature Cited Ambrose, H. W., III. 1969. A comparison of Microtus pennsylvanicus home ranges as determined by isotope and live trap methods. American Midland Naturalist 81: 535-555. Ambrose, H. W., III. 1973. An experimental study of some factors affecting the spatial and temporal activity of Microtus pennsylvanicus. Journal of Mammalogy 54: 79-110. Banks, E., R. Brooks, and J. Schnell. 1975. A radio- tracking study of home range and activity of the brown lemming (Lemmus trimucronatus). Journal of Mam- malogy 56: 888-901. Bendell, J. F. 1959. Food as a control of a population of white-footed mice, Peromyscus leucopus novebora- censis (Fischer). Canadian Journal of Zoology 37: 173-209. Brooks, R.J. and E.M. Banks. 1971. Radio-tracking study of lemming home range. Communications in Behavioral Biology 6: 1-S. Chute, F.S., W.A. Fuller, P. R. J. Harding, and T.B. Herman. 1974. Radiotracking of small mammals using a grid of overhead wire antennas. Canadian Journal of Zoology 52: 1481-1488. Drickamer, L. C. 1972. Experience and selection behavior in the food habits of Peromyscus: use of olfaction. Behaviour 41: 269-287. Drickamer, L.C. 1976. Hypotheses linking food habits and habitat selection in Peromyscus. Journal of Mam- malogy 57: 763-766. Dudley, D. 1974. Contribution of paternal care to the growth and development of the young in Peromyscus californicus. Behavioral Biology 11: 155-166. Eisenberg, J. F. 1968. Behavior patterns. /n Biology of Peromyscus (Rodentia). Edited by J. A. King. American Society of Mammalogists Special Publication Number 2. pp. 451-495. Graham, W. 1968. Daily activity patterns in the meadow vole, Microtus pennsylvanicus. Ph.D. dissertation, Uni- versity of Michigan, Ann Arbor, Michigan. 98 pp. Horner, B. E. 1954. Aboreal adaptations of Peromyscus, with special reference to the use of the tail. Con- tributions of the Laboratory of Vertebrate Biology of the University of Michigan 61: 1-84. Jackson, H.H.T. 1961. Mammals of Wisconsin. Uni- versity of Wisconsin Press, Madison, Wisconsin. 504 pp. Madison, D.M. 1977. Behavioral and _ sociochemical susceptibility of meadow voles (Microtus pennsylvanicus) to snake predators. American Midland Naturalist (in press). M’Closkey, R.T. 1975. Habitat dimensions of white- footed mice, Peromyscus leucopus. American Midland Naturalist 93: 158-167. MADISON: PEROMYSCUS MOVEMENTS AND HABITAT USE BY TELEMETRY 281 M’Closkey, R.T. 1976. Use of artificial microhabitats by white-footed mice, Peromyscus leucopus. American Mid- land Naturalist 96: 467-470. M’Closkey, R.T. and D.T. Lajoie. 1975. Determinants of local distribution and abundance in white-footed mice. Ecology 56: 467-472. Metzgar, L.H. 1971. Behavioral population regulation in the woodmouse, Peromyscus leucopus. American Midland Naturalist 86: 434-448. Millar, J.S. 1975. Tactics of energy partitioning in breed- ing Peromyscus. Canadian Journal of Zoology 53: 967-976. Mineau, P. and D. Madison. 1977. Radiotracking of Peromyscus leucopus. Canadian Journal of Zoology 55: 465-468. Myton, B. 1974. Utilization of space by leucopus and other small mammals. 277-290. Nicholson, A.J. 1941. The homes and social habits of the wood-mouse (Peromyscus leucopus noveboracensis) in southern Michigan. American Naturalist 25: 196-223. Pearson, P. G. 1959. Small mammals and old-field succes- sion on the piedmont of New Jersey. Ecology 40: 249-255. Shields, L. J. 1976. Telemetric determination of the activity of free-ranging rodents: The fine-structure of Microtus californicus activity patterns. Ph.D. dissertation, Uni- versity of California, Los Angeles, California. 95 pp. Synder, D. P. 1956. Survival rates, longevity, and popula- tion fluctuation in the white-footed mouse, Peromyscus leucopus, in southeastern Michigan. Miscellaneous Publi- cations of the Museum of Zoology of the University of Michigan 95: 1-33. Sokal, R.R. and F.J. Rohlf. 1969. Biometry. Freeman and Company, San Francisco. 776 pp. Stickel, L. F. 1968. Home range and travels. /n Biology of Peromyscus (Rodentia). Edited by J. A. King. Ameri- can Society of Mammalogists Special Publication Number 2. pp. 373-411. Stickel, L.F. and O. Warbach. 1960. Small-mammal populations of Maryland woodlot, 1949-1954. Ecology 41: 269-286. Terman, C. P. 1968. Population dynamics. /n Biology of Peromyscus (Rodentia). Edited by J. A. King. American Society of Mammalogists Special Publication Number 2. pp. 412-450. Whitaker, J. O., Jr. 1966. Food of Mus musculus, Pero- myscus maniculatis bairdi and Peromyscus leucopus in Vigo County, Indiana. Journal of Mammalogy 47: 473-486. Peromyscus Ecology 55: W.H. Received 5 November 1976 Accepted 13 July 1977 Prairie Fires and Pronghorn Use of Cactus JOHN G. STELFOX! and HAROLD G. VRIEND? 'Canadian Wildlife Service, Edmonton, Alberta 2Alberta Fish and Wildlife Division, Lethbridge, Alberta Stelfox, John G. and Harold G. Vriend. 1977. Prairie fires and pronghorn use of cactus. Canadian Field-Naturalist 91(3): 282-285. Abstract. Extensive prairie fires occur frequently on the Suffield Military Reserve as a result of lightning strikes and military operations. Pronghorns (Antilocapra americana) were observed feeding on prickly pear cactus (Opuntia polyacantha) growing on burnt-over areas. Fires had removed most spines from the cactus plants, providing a source of preferred forage that apparently attracted pronghorns from adjacent unburned ranges. The Suffield Military Reserve, a 2590-km? (1000-mi2) block of relatively undisturbed native prairie in southeastern Alberta (Figure 1), is owned by the Federal Government and managed Newe// by the Department of National Defence. Except for a 624-km2? (240-mi?) area of the Middle Sand Hills and land adjacent to the South Sas- katchewan River which has been used for Ralstone; ae LETHBRIDGE O kilometres Se ee o miles ®@ Raymond 40 @ Suffield MEDICINE HAT FIGURE |. Location of the Suffield Military Reserve in southeastern Alberta. 28 2 STELFOX AND VRIEND: 1977 FIRES AND PRONGHORN USE OF CACTUS 283 FIGURE 2. A small herd of pronghorns on a recent burn in the grassland. emergency grazing during periods of drought, the remainder has been used for military manoeuvres. In addition, gas exploration ac- tivities commenced in the northwest portion of the reserve in 1976. Extensive prairie fires are associated with lightning strikes and military operations. As much as 30% of the range is burned annually as a result of these fires (Cameron, unpublished report). It was on 4 November 1975, while conducting a range survey of the Suffield Block, that the authors recorded incidental observations of the prong- horn antelopes feeding and bedding on these burnt-over areas (Figure 2). Study Area The Suffield Military Reserve is located north of the city of Medicine Hat, adjacent to the South Saskatchewan River. It is generally described as mixed-grass prairie with undulating to rolling topography (Figure 2). Hilly areas and sand dunes are present within the Middle Sand mixed prairie range. Note the pattern of recent burns over Hills complex, and coulees are present near the river. The variable surficial deposits display dominantly brown Chernozemic soils. The grassland is predominantly a spear grass — blue grama community (Stipa comata — Bouteloua gracilis) as reported by Mitchell and Smoliak (1971) and by Jaques (1977). The incidence of silver sage (Artemisia cana) is low to moderate throughout most of the area. The pronghorn (Antilocapra americana) is the most abundant wild ungulate on the Reserve, numbering between 750 and 2000 animals depending on its seasonal: migratory patterns and weather conditions (Stelfox, unpublished report; Bibaud, unpublished report; Vriend SIA) Observations A close examination of the recently burned area on which the pronghorns were foraging revealed no new forage growth since the fire 284 THE CANADIAN FIELD-NATURALIST Vol. 91 oe eee Los FIGURE 3. A comparison of burnt versus unburnt cactus clones. The unburnt stem on the left was taken from a clone adjacent to the burn and placed next to a burnt and partially eaten clone. which occurred about one month previous. The only edible vegetation remaining was prickly pear cactus. The recent fire had removed most of the spines from the cactus plants (Figure 3) and the pronghorns had been actively pawing the cactus clones and foraging on the fleshy de- spined stems. Of about 30 cactus clumps observed within | hectare, all had been utilized. Close inspection of one large cactus clone approximately 1m in diameter, revealed 13 dislodged and 9 intact stems. It was estimated that over 50% of the green stems had been utilized in this burn. Discussion Food habitat studies of pronghorn antelopes in Alberta have revealed a low incidence of cactus in the diet. Mitchell and Smoliak (1971) recorded 1% mean volume of prickly pear cactus in the late fall diet of pronghorns in the Manyberries district, 128 km south of Suffield, and 7% and 12% mean volume of ball cactus (Mamillaria vivipara) in spring and fall diets, respectively, of pronghorns in the Newell area, 32 k west of Suffield. Twenty years of range— pronghorn relationships (data unavailable) in a 2.6-km? enclosure near Manyberries stocked with high densities of pronghorns suggested that cactus is more heavily utilized when other food sources are depleted (L. D. Gudmundson, Alberta Fish and Wildlife Division, personal communication). During the fall period, pronghorns tend to concentrate on areas having an abundance of green vegetation. They are commonly seen feeding on winter wheat (Triticum sativum) or alfalfa (Medicago falcata) fields in areas where this type of vegetation is available. The succulent nature of the vegetation promotes its use. Inasmuch as no cropland is available within the Suffield Reserve, the fall use of “spineless” cactus may constitute an important element of the pronghorn’s diet in this area. Management implications are undetermined, 1977 although it appears that restricted fires on areas with a prominence of cactus, especially in drought years, could provide a palatable, nutri- tious source of forage. A Colorado study (Schoop et al. 1977) determined through chemical analysis and micro-digestion trials that singed prickly pear was a palatable and nutritious food for livestock. Prickly pear contained 40% more soluble carbohydrates than alfalfa; however, it had a low (3.4%) digestible protein content. The digestibility of prickly pear was equal to or superior to that of high quality alfalfa hay. Literature Cited Jaques, D. 1977. The vegetation and effects of grazing STELFOX AND VRIEND: FIRES AND PRONGHORN USE OF CACTUS 285 on the eastern portion of the Suffield Military Reserve, Alberta. Jn Effects of livestock grazing on mixed prairie range and wildlife within PFRA pastures, Suffield Military Reserve. Edited by J.G. Stelfox. Canadian Wildlife Service Publication, Edmonton. Mitchell, G. J. and S. Smoliak. 1971. Pronghorn antelope characteristics and food habits in Alberta. Journal of Wildlife Management 35(2): 238-250. Schoop, M.C., E. J. Alford, and H. F. Mayland. 1977. Plains prickly pear is a good forage for cattle. Journal of Wildlife Management 30(1): 12-17. Vriend, H. 1977. Wildlife history. Jn Effects of livestock grazing on mixed prairie range and wildlife within PFRA pastures, Suffield Military Reserve. Edited by J.G. Stelfox. Canadian Wildlife Service Publication, Ed- monton. Received 22 November 1976 Accepted 23 April 1977 Notes Waterfowl Use of Exotic Wild Rice Habitat in Northern Saskatchewan DONALD G. PEDEN Canadian Wildlife Service, 115 Perimeter Road, Saskatchewan S7N 0X4 University of Saskatchewan Campus, Saskatoon, Peden, Donald G. 1977. Waterfowl use of exotic wild rice habitat in northern Saskatchewan. Canadian Field-Naturalist 91(3): 286-287. The density of waterfowl populations in the boreal forests of Saskatchewan is known to be much lower than that of the parklands and prairies to the south (Wellein et al. 1964; Anderson and Henny 1972). A few widely scattered pockets of relatively high density (Smith et al. 1964), however, occur in areas suchas the Saskatchewan River Delta (Dirschl and Goodman 1967). Surveys conducted during 1975 near La Ronge, Saskatchewan, suggest that planting of wild rice (Zizania aquatica) has created locally important high density waterfowl habitat. Wild rice is not native to Saskatchewan (Dore 1969), its nearest natural range being the east shore of Lake Winnipeg. Experimental seeding occurred in 1935 at Potato Lake, Pine Branch River, Upper Limestone Lake, and Nemeiben Lake near La Ronge (Neilson 1964). Verbal reports indicate that local residents also occasionally scattered seed in lakes and streams in a haphazard manner. In 1964, systematic seeding began (La Ronge Industries, personal com- munication) and presently La Ronge Industries, assisted by the Department of Northern Saskatche- wan, is attempting to expand the acreage of wild rice for commercial production. During the summer of 1975, the distribution of known stands of wild rice near La Ronge was mapped by canoe and by air photographs. Waterfowl were recorded both in the wild rice stands and in all non- rice areas between the stands. Counts were made during daylight hours. For each month, each site was visited once with the total number of birds seen having been recorded. The amount of non-rice habitat surveyed was not measured but it was certainly many times greater than the area of rice stands. Wild rice acreage was measured in | 1 locations near La Ronge. The total area (Figure 1) was approxi- mately 230 ha. This area excludes the Sikachu Lake rice field which may approach 75 ha and a few other small areas not exceding 4 ha each. Compared to a total area of more than 1,000,000 ha covered by the map in Figure |, the extent of wild rice is at present very small. Even though the total area of wild rice is small, the TABLE 1—Comparison of the total number of waterfowl (percent in parentheses) observed in wild rice and non-wild rice habitat during June, July, August, and October 1975 Rice Non-rice 1-13 June 183 (83) 37 (17) 1-18 July 525 (84) 102 (16) 29 July-15 August 168 (79) 45 (21) 11 October 1725 (99) 10 (1) Total 2601 (93) 194 (7) majority of the waterfowl numbers (Table 1), 93% of the 2795 sightings, occurred in wild rice habitat. Greatest waterfowl densities occurred during Oc- tober, suggesting use of wild rice as a staging area. Fortunately for those endeavoring to crop wild rice, this influx of birds occurred after the harvest. But evidence of damage to wild rice crops is noticeable (Kaz Parada, personal communication). Lesser Scaup (Aythya affinis), Goldeneye (Bucephala clangula), and Mallard (Anas platyrhynchos), respectively, ranked as the three most common species. Seventy- four of the 525 waterfowl in wild rice habitat during July were young of these three species indicating some use of wild rice habitat for breeding. The importance of these observations lies not in the magnitude and accuracy of the numbers but rather in the fact that waterfowl numbers were greatest in a very small exotic habitat type, namely wild rice. There are two plausible explanations for this phenomenon. Either waterfowl are attracted directly to the wild rice or wild rice is adapted to the same habitat features that waterfowl select. If the first explanation is true, then abundance of waterfowl has likely increased with man’s importation of wild rice. This trend may well continue if wild rice acreage continues to expand. If the second explanation is valid, one might expect that waterfowl, especially Lesser Scaup and Mallard, could bea useful indicator of potential sites for future wild rice seeding. 286 NOW 287 56° Z ~ MERIDIAN /7 s Nan. FIGURE |. Location of wild rice stands near La Ronge, Saskatchewan. The names and areas for each designated stand are as follows: 1, Mud Bay 5.8 ha; 2, East of Whitmore Island 3.4 ha; 3, North end of Nemeiben Lake 6.4 ha; 4, Three Portage Bay 16.7 ha; 5, Jackfish Bay 13.2 ha; 6, Miller channel 2.2 ha; 9, Pine Branch River 69.7 ha; 10 and 11, Howard Lake 11.0 ha; 12, Morning Lake 76.0 ha; 13, Trivet Lake 16.1 ha; and 16, Potato Lake 10.3 ha. Traces resulting from new seed were found at stands 7 and 8 (Head Lake) and 14 (Pisew Lake). Areas 17, 18, and 19 respectively, designate Egg Lake, the Sikachu River, and Waden Bay from which estimates of areas were not available. Whichever explanation is the case, this man-made alteration to what is often considered a pristine wilderness deserves future consideration. Acknowledgments Without the generous assistance of Kaz Parada, La Ronge Industries, and Oral Young, Economic De- velopment Branch, Department of Northern Sas- katchewan, this study would not have been possible. The author is also grateful for review and comments given by J. B. Gollop, G. D. Adams, and D. Lane, Canadian Wildlife Service. Literature Cited Anderson, D. R. and C. J. Henny. 1972. Population ecol- ogy of the Mallard. I: A review of previous studies and the distribution and migration from breeding areas. Bureau of Sport Fisheries and Wildlife, Washington, Reference Publication 105. 166 pp. Dirschl, J. H. and A. S. Goodman. 1967. Land capability for wildlife production and utilization in the western Saskatchewan River delta. Canadian Wildlife Service, Saskatoon. 233 pp. Dore, W.G. 1969. Wild rice. Canada Department of Agriculture, Research Branch Publication Number 1393. 84 pp. Neilson, J. D. 1964. Wild rice identification and propaga- tion in northern Saskatchewan. Saskatchewan Depart- ment of Agriculture, Prince Albert. 13 pp. Smith, R.H., F. Dufresne, and H. A. Hansen. 1964. Northern watersheds and deltas. Jn Waterfowl tomorrow. Edited by J. P. Linduska. United States Fish and Wild- life Service, Washington, D.C. pp. 51-66. Wellein, E. G. and H. G. Lumsden. 1964. Northern forests and tundra. /n Waterfowl tomorrow. Edited by J.P. Linduska. United States Fish and Wildlife Service, Washington, D.C. pp. 67-76. Received 4 November 1976 Accepted 5 March 1977 288 THE CANADIAN FIELD-NATURALIST Vol. 91 The Wolffish, cf. Anarhichas denticulatus, New to the Amundsen Gulf Area, Northwest Territories, and a Probable Prey of the Ringed Seal THOMAS G. SMITH Arctic Biological Station, Fisheries and Marine Service, Department of Fisheries and the Environment, P.O. Box 400, Ste-Anne-de-Bellevue, Quebec H9X 3L6 Smith, Thomas G. 1977. The Wolffish, cf. Anarhichas denticulatus, new to the Amundsen Gulf area, Northwest Territories, and a probable prey of the Ringed Seal. Canadian Field-Naturalist 91(3): 288. On 16 May 1974 while hunting Ringed Seals (Phoca hispida), two Inuit from Holman, Northwest Terri- tories, found the carcass of a Wolffish (Anarhichas sp.). The 13.6-kg, 127.0-cm-long carcass was found floating in the breathing hole beside a seal which they had shot on the ice. The hunting area was near Iluvilik Island (70°37’ N, 116°32’ W) in Prince Albert Sound on western Victoria Island. The area in the gill region and ventrally had been chewed away and the viscera were completely missing. The body showed many scratch marks similar to those inflicted by the claws of the foreflippers of the seal on Arctic Char fed to them in captivity. The fact that the Wolffish carcass was found in the breathing hole is strong evidence that the seal had been feeding on it. Few people in the Holman region had seen this species of fish before. A few older hunters knew the local Inuit name which is akoak or akoaksaluk (old woman fish). Two hunters, some years before, had found an even larger specimen frozen into the surface of the sea-ice near the same area, but did not know how the carcass had arrived there. Unfortunately, we were unable to collect the specimen or to make a positive species identification in the field. From photographs of the carcass, its gray color and large size, it has tentatively been identified as Anarhichas denticulatus Kroyer (D. McAllister, National Museum of Natural Sciences, personal communication). The closest known locality for A. denticulatus is 620 km to the north in Mould Bay, Prince Patrick Island (Walters 1953). Another closely related species, A. orientalis, is known from the Bering Sea and north Pacific Ocean (Barsukov 1959) and has been reported to the southeast of the present area in Bathurst Inlet (Legendre et al. 1975). The Ringed Seal which apparently successfully caught this specimen was shorter in total length (121 cm) than its prey. The Ringed Seal is not generally known as a predator of large fish and most data on its feeding habits come from the examination of stomach contents. From these it has been described as primarily a fish eater during the ice-covered months. Its main prey species is the Arctic Cod, Boreogadus saida, which rarely exceeds 160 mm in length and 35 gm in weight, in this area. It also feeds on the Saffron Cod, Eleginus sp., and on the Polar Cod, Arctogadus sp., which rarely exceed 400 mm and 500 mm in total length, respectively. Evidence of utilization of larger fish might easily be missed in analyses of stomach contents, especially when the head is not eaten. Identification of fish remains depends primarily on otoliths or a sufficiently large intact piece of the body. We know from observations on captured Ringed Seals that they can and will actively pursue and kill large fish in the confines of their holding pens. Observations of seals fed live Arctic Char, Sa/lvelinus alpinus, showed a consistent hunting behavior. The seal would actively pursue the fish, repeatedly biting them behind the head in the gill area. Large char took a fairly long time to subdue. A 7.71-kg char died after 22 min of pursuit and attack. We conclude that large fast-moving fish, such as Arctic Char, would not likely be a normal constituent of the Ringed Seal diet in the wild. The Wolffish, primarily a bottom feeder, however, is a rather sluggish swimmer which might easily be caught and eventually killed by a seal. Recent evidence indicating that the Ringed Seal is capable of deep and extended dives supports this idea (Geraci and Smith 1975). Literature Cited Barsukov, V.V.1959.Sem Zubatok (Anarhichadidae). Fauna SSSR Akademiia Nauk SSSR Zoologicheskii Institut 73: 1-171. Geraci, J. R.and T. G.Smith. 1975. Functional hematology of ringed seals (Phoca hispida) in the Canadian Arctic. Journal of the Fisheries Research Board of Canada 32(12): 2559-2564. Legendre, V.,J.G. Hunter, and D. McAllister. 1975. French, English and scientific names of marine fishes of arctic Canada. Syllogeus 7: 1-15. Walters, V. 1953. Notes on fishes from Prince Patrick and Ellesmere Islands, Canada. American Museum Novitates 1643. 17 pp. Received 22 February 1977 Accepted 6 May 1977 1977 NOTES 289 Locations of Winter Dens Utilized by Striped Skunks in Delta Marsh, Manitoba GRAHAM R. P. MUTCH Zoology Department, University of Manitoba, Winnipeg, Manitoba Present address: Interdisciplinary Systems Ltd., 528 St. James Street, Winnipeg, Manitoba Mutch, Graham R. P. 1977. Field-Naturalist 91(3): 289-291. R3T 2N2 R3G 3J4 Locations of winter dens utilized by Striped Skunks in Delta Marsh, Manitoba. Canadian Abstract. Locations of winter dens utilized by Striped Skunks in Delta Marsh, Manitoba were determined by tracking free- ranging skunks in snow. Skunks selected wet low-lying areas in preference to a drier sandy ridge and a dike (road embankment) for den sites. Higher soil temperatures, less extreme snow conditions, and greater substrate stability are possible explanations for den-site preference contrasting with the results of studies in other areas. Burrowing mammals that become inactive during winter must find or prepare suitable dens for occupancy during late fall, winter, and early spring. An optimal den site theoretically would be in stable well-drained soil in an area not subject to flooding in spring and would be sufficiently deep that the nest chamber would not freeze. This paper describes the locations of 19 dens used by Striped Skunks (Mephitis mephitis) during winter in Delta Marsh, Manitoba (50° 11’ N, 98°23’ W). This location is 5—6° of latitude south of the northernmost part of the skunk’s range at this longitude (Hall and Kelson 1959). Skunks undergo a near-complete cessation of surface activity in this area during winter (Mutch 1976). Data on soil temperatures and snow conditions in the various areas utilized are also presented. Study Area and Methods The study was conducted at the University of Manitoba Field Station, Delta Marsh on a 2.5-km strip of land adjacent to and paralleling Lake Manitoba. Based on differences in topography and flora, the area may be subdivided into three sub-areas. The first, a 50- to 200-m-wide sandy ridge immediately adjacent to the beach supports stands of Populus spp., Salix spp., Sambucus pubens, Acer negundo, and other tree and shrub species with a dense understory of herbs. In the second sub-area, a marsh-meadow community, Typha latifolia and Scirpus spp. fringe open water in natural channels and man-made borrow pits. Seasonally flooded sites support Scholochloa festucacea and Phragmites communis while drier meadows are characterized by Agropyron repens, Hordeum jubatum, Sonchus arvensis, Circium arvense, and Melilotus alba. The third sub-area is a 2- to 3-m-high dike embankment, the top of whichis a gravelled road. Its sides support a dense mixed stand of Bromus sp., Medicago sativa, and Melilotus alba, which may be mowed in summer. Walker (1959) has described the vegetation of Delta Marsh in detail. Weekly records of soil temperatures were obtained between late October and mid-May using telethermo- meter probes buried at a depth of 90 cm at one site in each sub-area described. The October marsh water table was at approximately 90 cm but in the ridge and dike embankment it was deeper. Snow depths were recorded weekly at each of 12 permanent snow stations in each sub-area, as well as an indicator of snow hardness, the pressure (in grams per square centimetre) required to compress the surface snow 2 or 3 cm or to break the surface crust. Occupied skunk dens were found in fall and spring by following skunk tracks in snow and seeking evidence of fresh digging together with skunk tracks at den mouths. Results Soil temperatures at a depth of 90 cm in each sub- area are depicted in Figure |. Prior to heavy snowfall in autumn, rapid declines in soil temperature occurred at all three locations. After the arrival of heavy snow, marsh soil cooled very gradually. Soil temperatures at a depth of 90 cm remained above 1°C for the rest of the winter. In May, warming of marsh soil was also very gradual. Changes in temperature of soil at the same depth in the wooded ridge followed a similar pattern, although soil here was consistently about 1°C cooler than in the marsh (Figure 1). The coldest soil was in the dike embankment where temperatures as low as —3°C were recorded in mid- to late February. Snow depths varied considerably in different sub- areas (Figure 2). The 130-cm cumulative snowfall recorded at the University Field Station was subject to relatively little drifting in the marsh where vegetation tended to hold it in place. Because little compaction occurred here the snow that accumulated remained soft and loose, generally requiring pressure of less than 6 g/cm? to compact it 2 or 3 cm. The dike was 290 10 @ C @—— @ MARSH 8 eo O——O DIKE 8 AN \ \ Vv VY RIDGE N pel V7. "\ a % \e O r S ole op) al “A z Ve = eh \ -@ e@ < ee ) Ga 0 iS \ e y ss 7 \ Vis ©». ee [ SP \ /s ry = YS Ve ‘e Peg ee: Ty ne \ AD \ = / \ Te Ys Sy oye OR wos. AVA WE (e) ) Nv / op © \ hele (0) N D J F M A M MONTHS (1974-1975) FIGURE |. Weekly soil temperature readings at 90-cm depth in the marsh, dike embankment, and wooded ridge sub-areas at Delta Marsh, Manitoba for the period October 1974 to May 1975. The arrow indicates the first heavy snowfall of the season. windswept and the road surface snow-free. Mowed embankment slopes trapped little snow, which melted in early March, especially on the east-facing slope. Snow rarely became hard here. On the wooded ridge snow blown from the adjacent lake accumulated into deep hard drifts. These drifts were deepest between the center of the ridge and the lake. Drifts were frequently very hard, requiring up to 100 g/cm2 to compact the snow 2 or 3cm. In areas where the ridge was more than 100m wide, drifts were often absent on the downwind (marsh) side. Few skunk tracks were found in November and December and none were found in January and February. Tracks became common in March and April. Skunk tracks on the ridge were always uncommon except immediately adjacent to the marsh. Greatest concentrations of tracks were found in the marsh. I found 19 dens that were repeatedly utilized by skunks and none of these was located in the ridge. Three were on the marsh-ridge boundary where the flora was similar to that of the marsh and soil was similar in wetness and structure to marsh soil. These THE CANADIAN FIELD-NATURALIST Vol. 91 sites were beyond the range of deep hard drifts. No active skunk den was found on the dike embankment. Three were found on other, slightly raised road embankments and one beneath an old culvert. Snow in these areas was less than 0.75 m deep, and soft. Vegetation consisted of forbs and grasses. In no case were these dens in snow conditions similar to those prevailing in the wooded ridge. The remaining 12 dens were located in the marsh, eight in stands of Phragmites communis. \t was not possible to deter- mine what species had originally excavated the dens although the marsh dens showed signs of having been dug by skunks. Discussion This survey presents evidence that Striped Skunks in Delta Marsh occupy winter dens located in flat low- lying areas, often in Phragmites stands. Although surface litter may be dry, these sites have heavy, a—a Cumulative Snowfal) 140 e—e Marsh o—cC Dike ACA y—v Ridge aca’ SNOW DEPTH (em) MONTHS(1974-1975) FIGURE 2. Weekly mean snow depths in the marsh, on the dike embankment, and in the wooded ridge (12 stations in each sub-area) at Delta Marsh, Manitoba, for the period October 1974 to May 1975. Cumulative snowfall was determined by summing records for all snowfall measured at the meteorological station at the main University Field Station compound in Delta Marsh. IY poorly drained soils and a high water table. Complete inundation invariably occurs in March or early April upon melting of the snow. These features are apparently disadvantageous to winter denning in the marsh sub-area. Burrows dug by hibernating rodents are present in the ridge, and skunks could possibly modify these to their own purposes if denning in the ridge were otherwise acceptable. Other studies in- dicate that burrowing mammals consistently avoid wet areas for winter den sites. Skunks have been found to select hilly or rolling country for their winter dens, which are often on well-drained slopes (Selko 1938: Scott and Selko 1939; Allen and Shapton 1942; Verts 1967). Bailey (1971) found that skunks in a Lake Erie marsh denned exclusively in artificial dikes. House- knecht (1969) reported that skunks denned almost entirely in upland sites in winter; in summer a significant proportion of nest sites, many of them aboveground, were reported in lowland areas by Houseknecht (1969) and Storm (1972). Local conditions peculiar to the study area may necessitate winter denning in sites avoided in other environments. True uplands are lacking in the strip of marsh and wooded ridge paralleling Lake Manitoba. The wooded ridge, despite dryer soils, a lower water table, and very localized spring flooding has limita- tions. The soil is essentially pure sand and may be too unstable to support construction of large dens, although hibernating rodents do den there. Hard snow, 2 m or more in depth, blankets the ridge until May and could impede or even prevent entrance to, or emergence from, dens in spring. Sunquist (1974) believed that particularly deep and heavy snow was at least partially responsible for delaying spring emer- gence of skunks in Minnesota. Easy entrance to the den is essential during the rut in March and April (Verts 1967). The dike embankment is well-drained and its soil is firm. It is blown clear of snow, and soil cemperatures at 90 cm depth fell to below freezing for extensive periods. Lower ambient temperatures at more northern latitudes may make it necessary that skunks den in the warmest areas possible. Soft snow provides insulation to soil in the marsh and is not deep enough to trap skunks in their dens. The air layer between the soil and the previous summer's Phrag- mites growth, which has been flattened by the weight of the snow, is excellent insulation. Ground water in wet areas is a reservoir of heat ensuring higher soil temperatures (the recorded minimum marsh sub-area temperature at 90 cm depth was 1.1°C, compared to 0.1 and -3.0°C in the ridge and dike respectively). These factors are believed to be important in winter den-site selection by Striped Skunks in Delta Marsh. This marsh is excellent habitat for skunks, partic- ularly in relation to seasonal food abundance, in- cluding voles, duck eggs, and frogs. The absence of NOTES 291 sites preferred for denning in other regions has not prevented skunks in Delta Marsh from reaching high populations, estimated at 20-32/km? by Lynch (1972). It is therefore evident that the Striped Skunk has adapted to the local conditions and that marsh dens do in fact fulfill their den requirements. Acknowledgments This study was financed by the National Research Council of Canada (through grants to M. Aleksiuk, University of Manitoba, my M.Sc. supervisor), Manitoba Department of Mines, Resources and Environmental Management (to the University of Manitoba Field Station), and Gulf Oil Canada Limited (to the author). Personal maintenance funds were received from the same sources. Dr. Aleksiuk’s assistance in all phases of this study is greatly appreciated. This note is Publication Number 43 resulting from research done at the University of Manitoba Field Station (Delta Marsh). Literature Cited Allen, D. L. and W.W. Shapton. 1942. An ecological study of winter dens, with special reference to the eastern skunk. Ecology 23: 59-68. Bailey, T. N. 1971. Biology of striped skunks on a south- western Lake Erie marsh. American Midland Naturalist 85: 196-207. Hall, E. R.and K. R. Kelson. 1959. The mammals of North America. Ronald Press, New York. Houseknecht, C. R. 1969. Denning habits of the stnped skunk and the exposure potential for disease. Bulletin of the Wildlife Disease Association 5: 302-306. Lynch, G. M. 1972. Effects of strychnine control on nest predators of dabbling ducks. Journal of Wildlife Manage- ment 36: 436-440. Mutch, G. R. P. 1976. Aspects of winter dormancy in the striped skunk, Mephitis mephitis. M.Sc. thesis, University of Manitoba, Winnipeg, Manitoba. vill + 89 pp. Scott, T. G. and L. F. Selko. 1939. A census of red foxes and striped skunks in Clay and Boone Counties, Iowa. Journal of Wildlife Management 3: 92-98. Selko, L. F. 1938. Notes on the den ecology of the striped skunk in Iowa. American Midland Naturalist 20: 453-463. Storm, G. L. 1972. Daytime retreats and movements of skunks on farmlands in Illinois. Journal of Wildlife Management 36: 31-45. Sunquist, M. E. 1974. Winter activity of striped skunks (Mephitis mephitis) in east-central Minnesota. American Midland Naturalist 92: 434-446. Verts, B. J. 1967. The biology of the striped skunk. Univer- sity of Illinois Press, Urbana, Illinois. vii + 218 pp. Walker, J. M. 1959. Vegetation studies on the Delta Marsh, Delta, Manitoba. M.Sc. thesis, University of Manitoba, Winnipeg, Manitoba. xii + 203 pp. Received 19 June 1976 Accepted 27 March 1977 292 THE CANADIAN FIELD-NATURALIST Vol. 91 Age and Fecundity of the Tadpole Madtom, Noturus gyrinus, on Long Point, Lake Erie ROBIN MAHON Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1 Mahon, Robin. 1977. Age and fecundity of the Tadpole Madtom, Noturus gyrinus, on Long Point, Lake Erie. Canadian Field-Naturalist 91(3): 292-294. Abstract. Specimens collected from lagoons on Long Point, Lake Erie, in May and August, belonged to ages 1+, 2+, and 3+, as determined from vertebrae. The relationship of Log Fecundity = 1.918 log SL -0.148 log EW -1.750, where SL is standard length and EW is mean weight of one egg, was found for 33 ripe females collected in May. There is no published study of the biology of the Tadpole Madtom, Noturus gyrinus, in Canada (Scott and Crossman 1973) and little elsewhere (Carlander 1969). This species has been aged using vertebrae (Hooper 1949), and an average number of 50 eggs per female, with a maximum of 93 has been reported (Everman and Clark 1920). Scott and Crossman (1973) noted a specimen with 117 eggs. Methods Specimens were collected from lagoons on the eastern end of Long Point, Lake Erie (42°32’N, 80°07’ W), using an electric fish shocker or after treatment with rotenone (Mahon and Balon 1977). Sample 1, taken in the last week of May 1975, was from the inshore of a large lagoon (about 25 ha) in depths of less than | m. Sample 2, taken in the first week of August, was from several small, isolated lagoons (up to 600 m?) peripheral to the same large lagoon. After preservation in 10% formalin, fish were sexed, weighed, and their standard and total lengths measured to the nearest millimetre. Ovaries con- taining ripe eggs were removed and the number and combined weight of these eggs recorded. Division of the combined weight of ripe eggs by their number gave the mean weight of one ripe egg (EW) for each female. Two fish from each |-mm-length interval in each sample were aged using vertebrae. A vertical incision was made immediately posterior to the dorsal fin, and four vertebrae anterior to the incision removed. After being allowed to dry, the vertebrae were separated using a thin blade, and the surfaces of the centra were viewed under direct light using a binocular micro- scope. Dark and light growth bands were visible on both surfaces of the centra. The annulus was considered as a transition point from a dark band toa light band. The age assigned each fish was the same as the number of annuli observed, with a plus sign to indicate growth subsequent to annulus formation (Balon 1972). Results Length frequency histograms for samples | (May) and 2 (August) are shown in Figure |. Table | shows mean standard length, range in length, and the number of males and females at each age for each sample. Ages are also superposed on Figure | so that age groups as obtained from the vertebrae may be compared to the peaks of the histograms. The relationship between weight (W) and standard length (SL) for all fish was best expressed by a linear regression of logarithmic transformed data. Two such regressions were fitted, one for the ripe females: log W = 3.18 log SL — 4.86 (r = 0.97), and the second for all other fish: log W=2.91 log SL -—4.533 (r=0.97). The regression coefficients for these regressions are significantly different (P<0.05). TABLE !—Mean standard length (mm), range of length, and number of males and females at each age in May and August, for Noturus gyrinus Sample | (May) Sample 2 (Aug.) ]+ 2+ 3+ ]+ 2+ 3+ Mean length 36 55 63 52 75 83 Range DSS) 45-65 60-69 40-66 58-85 80-87 Number of males 128 32 9 30 58 6 Number of females 129 25 2 32 22 l 1977 NOTES 293 1+ 20 —_o——————_ August acu 3+ 10 mean > UO Z 5 30 eT SEIN ESL ©) 2+ Lu ee ri 20 pa May 10 0) 10 20 30 40 50 60 70 80 90 STANDARD LENGTH (mm) Ficure |. Length-frequency histograms for the Tadpole Madtom, Noturus gyrinus, in May (325 individuals) and August (149). Horizontal lines give the range in length at each age with the mean length indicated by a closed circle. Standard length and total length (TL) are related in the following way: log TL = 0.958 + 0.152 log SL, as determined by a linear regression of log TL onlog SL. Ripe females were taken in both May and August. Those from August showed signs of having already spawned some of their eggs and were not used for fecundity counts. In May, 10% of age 1+ females, 72% of age 2+ females, and both age 3+ females were mature. The number of ripe eggs per female varied from 43 to 160. Only 65.1% of this variation can be accounted for by variation in standard length whena linear regression of log F on log SL is fitted. A further 6.2% can be accounted for by considering variation in egg weight, as was done by Scott (1962). The regression equation for the trivariate relation- ship of log F on log SL and log EW is log F=1.918 log SL-0.148 log EW -1.750 (r = 0.844). There is significant positive correlation between log SL and log EW (r = 0.703). Discussion The absence of 0+ fish in the August sample is probably owing to their small size at this time. Case (1970) reports that 0+ Noturus gyrinus emerged in early August in a Manitoba river. For sample | fish, mean lengths at ages 1+ and 2+ correspond to peaks in the length-frequency histogram (Figure 1), suggesting that this method of aging is valid. There is no distinct peak of 3+ fish, however, and the conclusion that individuals do survive to 3+ years rests on the vertebral growth ring counts alone. The only comparable growth study is that of Hooper (1949) for Noturus gyrinus from Demming Lake, Minnesota, where growth was faster than in Long Point lagoons. In Demming Lake, where this species was dominant in standing crop and numbers, mean standard lengths of 26 mm (range 15-35 mm), 62mm (range 43-85mm), and 89mm (range 78-104 mm) had been achieved by age 0+, I+, and 2+ fish respectively on August 15. Though growth was faster in Demming Lake than on Long Point, mortality between ages 1+ and 2+ appears to have been much higher (97.4% and 77.8% respectively). Had mortality of age 2+ fish in Demming Lake been similarly high there would have been only seven or eight 3+ fish in the whole lake from which sub-samples were taken for aging. Considering the percentage mortality between ages I+ and 2+ in Long Point lagoons the finding of individuals aged 3+ is not unexpected. Noturus gyrinus nests and guards a few large yolky eggs which are usually hidden (Hankinson 1908; Bailey 1938). For this reproductive style, longevity and the accompanying increase in size and experience 294 would seem advantageous, yet most females mature at age 2+ and therefore probably reproduce only once. Case (1970) found that toxin injected directly from the pectoral spines of live Noturus gyrinus into the lateral musculature of a sauger, Stizostedion canadense, a northern pike, Esox /ucius, and a rainbow trout (fry), Salmo gairdneri, produced varying degrees of immobilization almost immediately. Esox Jlucius (30 cm) was most severely affected, turning belly up after 15 min. The manufacture of a toxin could serve as a substitute for increased size (Hairston et al. 1970) enabling Noturus gyrinus to defend its nest effectively against large predators either actively or as the result of learned avoidance on the part of the predator, and at the same time retain the small size needed for exploitation of its accustomed habitat. Acknowledgments I am grateful to Michael Chadwick and Judy Peat for their assistance in the field. David Noakes criticized the original manuscript, and Jan Hines offered valuable advice on the statistical treatments. The study was supported by a National Research Council of Canada operating grant to Eugene Balon, University of Guelph. Literature Cited Bailey, R M. 1938. The fishes of the Merrimack watershed. Jn Biological survey of the Merrimack watershed. New Hampshire Fish and Game Department Survey Report 3. pp. 149-185. Balon, E. K. 1972. Possible fish stock size assessment and available production survey as developed on Lake Kariba. THE CANADIAN FIELD-NATURALIST Vol. 91 African Journal of Tropical Hydrobiology and Fish- eries 2: 45-73. Carlander, K. D. 1969. Handbook of freshwater fishery biology. Volume |. Life history data on freshwater fishes of the United States and Canada, exclusive of the Perci- formes. Iowa State University Press, Ames, Iowa. Case, B.E. 1970. An ecological study of the tadpole madtom, Norurus gyrinus, (Mitchell): with special refer- ence to movements and population fluctuations. M.Sc. thesis, University of Manitoba, Winnipeg, Manitoba. Everman, B. W. and H.W. Clark. 1920. Lake Maxin- kuckee: a physical and biological survey. Indiana Depart- ment of Conservation Publication |: 1-660. Hairston, N. G., D. W. Tinkle, and H. M. Wilbur. 1970. Natural selection and the parameters of population growth. Journal of Wildlife Management 34: 681-690. Hankinson, T.L. 1908. A biological survey of Walnut Lake, Michigan. Report of the State Biological Survey of Michigan 1907: 153-288. Hooper, F. F. 1949. Age analysis of a population of the ameiurid fish, Schilbeodes mollis (Hermann). Copeia 1949: 34-38. Mahon, R. and E.K. Balon. 1977. Fish community structure in lakeshore lagoons on Long Point, Lake Erie, Canada. Environmental Biology of Fishes 2. (In press.) Scott, D. P. 1962. Effect of food quantity on fecundity of rainbow trout, Sa/mo gairdneri. Journal of the Fisheries Research Board of Canada 19: 715-730. Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin 184. Received 4 October 1976 Accepted 11 May 1977 The Little Gull (Larus minutus) in Arctic North America STEPHEN R. JOHNSON and WILLIAM J. ADAMS LGL Limited, Environmental Research Associates, 10110 124th Street, Edmonton, Alberta T5N 1P6 Johnson, Stephen R. and William J. Adams. 1977. The Little Gull (Larus minutus) in Arctic North America. Canadian Field-Naturalist 91(3): 294-296. On 16 June 1975, at approximately 1830 hours Yukon Standard Time (YST), we observed a Little Gull (Larus minutus) in association with approxi- mately 30 Arctic Terns (Sterna paradisaea) man- oeuvering over a flooded tundra marsh approxi- mately | km southwest of the Komakuk Beach DEW Station, Yukon Territory (69°36’N, 140°11’ W). WJA first watched the gull from a distance of 20-30 m for approximately 10-15 min with the aid of 8 X 35 binoculars and made his tentative identi- fication froma field guide (Robbins et al. 1966) that he carried with him. Approximately 10 min later, we both observed a Little Gull at approximately the same location; we again compared the bird under scrutiny with the painting and description in the field guide. Neither of us had seen a Little Gull previously. Although the bird was silent, we recorded the details of its behavior and appearance. The gull was hooded and was obviously smaller than the terns with which it was associated. The undersides of its wings were slatey-black and the uppersides of the wings were dark gray; we noted no other markings on the wings except their white posterior and distal margins. The gull had red legs and a red bill. We saw Sabine’s 1977 Gulls (Xema sabini) in the Komakuk area previous to this sighting of the Little Gull, and we were careful to look for characteristics that distinguished this new gull from a Sabine’s Gull. We continued surveying the area near this marsh during the evening of 16 June and at approximately 2000 hours YST we found two Little Gulls there, also associated with Arctic Terns. In the afternoon of 17 June, we saw two Little Gulls flying northwest over a portion of the still frozen Beaufort Sea approxi- mately 300 m from the beach at Komakuk. Both of these birds were heard giving their “kek-kek-kek” call; they were not associated with any other species. The first North American specimen of a Little Gull was collected during the first Franklin Expedition sometime between 1819 and 1822, apparently in the area between the Coppermine River, Northwest Territories, Canada and York Factory, Manitoba, Canada (Baillie 1963, pp. 95-97). Baillie (1963) recorded the apparent invasion of eastern North America by the Little Gull; he documented several Little Gull specimens taken in eastern North America since 1887 and the discovery reported by Scott (1963) of the species’ first nest and eggs in the New World on 1 June 1962 near Oshawa, Ontario. Tozer and Richards (1974, pp. 342-346) described subsequent nesting activities in the Oshawa area and near Rondeau Park and Cranberry Marsh, Ontario. Erdman and Steffen (in Tessen 1975) reported Little Gulls nesting in north-central Wisconsin during the summer of 1975, the first nesting record of this species in the United States. The Little Gull is now regularly sighted and is fairly common in these portions of the Great Lakes region. The first recorded appearance of the Little Gull along the west coast of North America was on 16 November 1968 near Riverside, California (McCaskie 1969) and during recent years Little Gulls have appeared regularly along the southwestern coast of British Columbia (Tatum 1973; Campbell et al. 1974; M. G. Shepard, personal communication). MeNicholl (1974) recently reported a possible sighting of a Little Gull in southern Manitoba, and Switzer (1974) reported a recent observation of one in Regina, Saskatchewan. Green (1974) reported Little Gulls associated with Arctic Terns in Duluth, Minnesota. In the north, the only comparatively recent New World specimen of a Little Gull was a male collected by Nero (1963) on 26 June 1963, 11 km(7 mi) south of Beaver Point (59°2’ N, 109° W) on the south side of Lake Athabasca, Saskatchewan. More recently, Pittaway and Nero (1971) reported a sighting of this species at Churchill, Manitoba. Baillie (1969) mentioned that in recent times the NOTES 295 Little Gull has been a rare but regular visitor in eastern North America and that this species is frequently seen during winter, fall, and spring in company with Bonaparte’s Gulls (Larus philadelphia). Baillie (1951) hypothesized that Little Gulls “colonized” North America from Siberia. According to this hypothesis Little Gulls may have migrated southeast from their Siberian nesting area to the Bering Sea where they may have encountered breeding Bonaparte’s Gulls (Baillie 1951). After this encounter, according to Baillie, both species then presumably continued southeast to the eastern wintering grounds of Bonaparte’s Gull along the North American east coast. Our coastal Yukon observations, viewed in light of the specimen from Lake Athabasca and the initial Franklin Expedition specimen, lend superficial cre- dence to Baillie’s hypothesis. Neither Dement’ev et al. (1951) nor Vaurie (1965), however, describe nesting areas of the Little Gull in far eastern Siberia or in the vicinity of the Bering Sea, although Dement’ev et al. (1951) do mention the possibility of Little Gulls wintering in the Sea of Okhotsk. Further, Bonaparte’s Gulls nest in only a few isolated places near the Bering Sea coast in western Alaska and there are no records of Little Gull specimens or sightings in Alaska (Gabrielson and Lincoln 1951; D. D. Gibson, per- sonal communication). The probability seems low that an encounter between these two species would occur in the Bering Sea area. In the Old World, Little Gulls commonly winter in large numbers along the coast of Great Britain and along the Iberian and Mediterranean coasts (Vaurie 1965; Brunn 1970). Thus it seems more probable that Little Gulls originally “colonized” eastern North America from their wintering grounds in Europe. Subsequently, through their association with Bona- parte’s Gulls, as suggested by Baillie (1951) and as indicated by more recent sightings (Pittaway and Nero 1971; McNicholl 1974; and others), Little Gulls may have wandered to the Canadian Arctic from their wintering grounds in eastern North America. Acknowledgments We thank W.W.H. Gunn, W. J. Richardson, R. A. Davis, J. G. Ward, D. V. Weseloh, and M. K. McNicholl for their comments on this paper. G. F. Searing, D. D. Gibson, C. E. Tull, and W. E. Renaud read an earlier draft and made helpful comments. O. L. Austin, Jr., also made helpful suggestions. The Canadian Wildlife Service provided funds (through the Canadian Beaufort Sea Project) that enabled us to study sea birds along the Beaufort Sea coast. We thank the Canadian Wildlife Service for permitting us to publish our observations. 296 THE CANADIAN FIELD-NATURALIST Literature Cited Baillie, J. L. 1951. Ontario-western New York Region. Audubon Field Notes 5(5): 286-287. Baillie, J. L. 1963. Three bird immigrants from the Old World. Transactions of the Royal Canadian Institute (Part 2) 34: 95-105. Baillie, J. L. 1969. Birds in Ontario—western invaders. Ontario Naturalist 7(1): 28-30. Brunn, B. 1970. Birds of Europe. McGraw-Hill Book Company, New York. 319 pp. Campbell, R. W., M. G. Shepard, B. A. MacDonald, and W. C. Weber. 1974. Vancouver birds in 1972. Vancouver Natural History Society, Vancouver, British Columbia. 96 pp. Dement’ev, G. P., N. A. Gladkov, and E. P. Spangenberg. 1951. Birds of the Soviet Union. Volume III. Edited by G. P. Dement’ev and N. A. Gladkov. Gosudorstuennoe Izdolel’stvo “Sovietskaya nauka,” Moskva. (Translated from Russian by Israel Programs for Scientific Trans- lations, Jerusaleum, 1969.) 1121 pp. Gabrielson, I. N. and F.C. Lincoln. 1951. The birds of Alaska. Stockpole Company, Harrisburg, Pennsylvania and Wildlife Management Institute, Washington, D.C. 922 pp. Green, J.C. 1974. Arctic Tern and Little Gull as migrants in Duluth. Loon 46 (2): 52-57. McCaskie, G. 1969. Fall migration, August 16, 1968 — November 30, 1968. Regional Report of the Southern Pacific Coast Region. Audubon Field Notes 23(1): 106-112. Vol. 91 MeNicholl, M. K. 1974. A probable Little Gull record for southern Manitoba. Blue Jay 32(4): 229-230. Nero, R. W. 1963. Birds of the Lake Athabasca Region, Saskatchewan. Saskatchewan Natural History Society, Special Publication 5. 143 pp. Pittaway, R. and R. W. Nero. 1971. Recent bird notes of interest for Churchill, Manitoba. Blue Jay 29(2): 60-63. Robbins, C. S., B. Brunn, and H. S. Zim. 1966. A guide to field identification—Birds of North America. Golden Press, New York. 240 pp. Scott, G. A. 1963. First nesting of the Little Gull (Larus minutus) in Ontario and in the New World. Auk 80(4): 548-549. Switzer, F. 1974. Little Gull visits Regina. 32(1): 46-48. Tatum, J.B. 1973. Annual bird report for Southern Vancouver Island — 1972. Victoria Natural History Society, Victoria, British Columbia. 80 pp. Tessen, D. D. 1975. The nesting season: June 1—July 31, 1975. Western Great Lakes Region. American Birds 29(5): 974-978. Tozer, R.G. and J.M. Richards. 1974. Birds of the Oshawa-Lake Scugog Region, Ontario. Tozer and Richards, Oshawa, Ontario. 384 pp. Vaurie, C. 1965. Birds of the Palearctic Fauna. Volume II, Nonpasseriformes. H. F. and G. Witherby Ltd., London. 763 pp. Blue Jay Received 5 October 1976 Accepted 21 February 1977 Summer Food Habits of Golden Eagles in Southwestern Alberta D. A. BOAG Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9 Boag, D. A. 1977. Summer food habits of Golden Eagles in southwestern Alberta. Canadian Field-Naturalist 91(3): 296-298. Olendorff (1976), in reviewing the literature on the food habits of Golden Eagles (Aquila chrysaetos) in North America, requested that “others... report their findings to give us a still better assessment of the economic impact of Golden Eagles.” This note records the food habits of a pair of Golden Eagles during the summers of 1955 and 1958 at a nest site in southwestern Alberta. At that time the eagles occupied a nest on a cliff face of Missing Link Mountain (114°38’ W, 50°39’N), 30km west of Turner Valley, Alberta. | Periodic visits to the nest site spanned a period of 106 days in 1955 (5 May to 18 August when the single eaglet fledged) but only 58 days in 1958 (2 June to 29 July during the nestling period of the single eaglet). I was able to record only the unconsumed remains of prey present in the nest on each visit. Nearly all bird and mammal remains had at least one limb intact, enabling me to mark them by removing the toes; thus, I could leave them at the nest site as potential food, yet avoid recording them more than once. In each instance I was able to identify the species of prey and, in most cases, to determine their sex and age. The nest site was visited on 79 days in 1955 (18 in May and 3 in June during incubation, and 21 days in June, 25 in July, and 12 in August during the nestling period) and on 11 days in 1958 (5 in June and 6 in July during the nestling period). The prey items are recorded in Table 1. In both years, mammals made up more than 80% of the diet; the primary prey species was the Columbian ground squirrel (Spermophilus columbianus). This 1977 NOTES 207) TABLE 1—Prey items recorded at a Golden Eagle nest site located in southwestern Alberta. Data were recorded during the nestling periods of 1955 and 1958 Percent of total, 1955! Prey species Individuals Birds Galliformes Dendragapus obscurus 8 Bonasa umbellus I Piciformes Colaptes auratus 0 Passeriformes Pica pica 0 Total 9 Mammals Carnivora Mustela frenata 1 Rodentia Spermophilus columbianus 80 S. lateralis 4 Thomomys talpoides l Neotoma cineria l Lagomorpha Lepus americanus I Artiodactyla Odocoileus hemionus 2 Total 90 Percent of total, 1958 Biomass? Individuals Biomass? 15 8 12 <1 3 2 0 3 <1 0 3 <1 15 17 15 <1 0 0 77 79 82 D 5 3 (Nyl.) Arn. (C) #15 and #17. Caloplaca sp. (O) #5. This saxicolour lichen from the upper littoral zone has a dark gray thallus and dark orange apothecial discs with dark gray margins. It is common in the same community all along the rocky coast of the Queen Charlotte Islands (British Colum- bia) and must be widespread. It resembles C. cerina (Ehrh.) Th. Fr. var. chlorina (Flot.) Mull. Arg. but has a smoother thallus and different ecology (exposed littoral vs. shaded moist rocks); however, it may bea morphotype of this taxon. Caloplaca cerina var. chlorina has not been reported from British Columbia. *Caloplaca thallincola (Wedd.) Du Rietz (O) #6. This lobate Caloplaca is quite common in the British Isles where it is also a maritime species associated with Verrucaria maura (Duncan 1970). It isaspecies of the north- and west-European coasts (Poelt 1969). Lecanora sp.(O) #11. This material appears to be in the Lecanora subfusca group close to L. campestris (Schaer.) Hue morphologically, but its C+ orange thallus cortex (probably containing a xanthone) more closely ties it to L. fugiens Nyl. with regard to chemistry. Parmelia saxatilis (L.) Ach. (A) #1. ** Spilonema revertens Nyl. (O) #7 and 4. As in Europe, this species apparently is widely distributed in the north, but largely overlooked (see Poelt 1969). It has been reported by Ahti (1964) from a lakeshore in the southern boreal zone of Ontario, and by Thomson et al. (1969) from an open black spruce forest near Great Slave Lake (Northwest Territories). Sticta weigelii (Ach.) Vain. (O) #2. Verrucaria maura Wahlenb. ex Ach. (A) #9 and 3. occasional, C = common, Site 2. Exposed rocky point north of marine station. (a) Rocky Shore Acarospora fuscata (Schrad.) Arn. (O) #28. Aspicilia cinerea (L.) Korb. (O) #38. Buellia stellulata (Tayl. in Mack.) Mudd (O) #34. Caloplaca sp. (O) (See comments above) #30. Caloplaca marina (Wedd.) Zahlbr. (O) #39. Caloplaca scopularis (Nyl.) Lett. (O) #37. 5Harris (1975) points out that this species is actually not an Arthopyrenia. Until the new combination made by Harris in his thesis is validated, we are leaving the name unchanged. 308 Lecanora muralis (Schreb.) Rabenh. (O) #33. *Ochrolechia androgyna (Hoffm.) Arn. var. saxorum (Oeder) Vers. (O) #35. Parmelia stictica (Del.) Nyl. (C) #23. Physcia adscendens (Th. Fr.) Oliv. (O) #36. Placopsis gelida (L.) Linds. (C) #27. Rhizocarpon disporum (Naegeli ex Hepp) Mull. Arg. (C) #26. Spilonema revertens Nyl. (O) #29. Sticta weigelii (Isert ex Ach.) Vain. (O) #22. Verrucaria maura Wahlenb. ex Ach. (A) #41. Xanthoria candelaria (L.) Th. Fr. (O) #32. (b) Shady Rockface on Overhanging Cliff Porina chlorotica (Ach.) Mull. Arg. (A) #42. This maritime population on the British Columbia coast differs from European material of the species and may represent a new taxon (Harris 1975). The situation is discussed more fully by Brodo (1976). Verrucaria maura Wahlenb. ex Ach. (O) #44. (c) Low Shrub above High Tide Dimerella lutea (Dicks.) Trev. (R) #47 and 55. Lecania sp. (R) #56. This Lecania seems close to L. nylanderiana Mass. but differs from that species in having one-septate, constricted spores, larger apo- thecia, and persistent, thick, thalline margins. The thallus and apothecial discs are C-. Lepraria incana (L.) Ach. (O) #58 and 64. Leptogium palmatum (Huds.) Mont. (O) #51. Lobaria pulmonaria (L.) Hoffm. (C) #48. Nephroma laevigatum Ach. (O) #45. Pannaria microphylla (Sw.) Mass. (O) #46. Peltigera aphthosa (L.) Willd. (O) #57. Peltigera membranacea (Ach.) Nyl. (A) #49. Peltigera venosa (L.) Baumg. (O) #52. Proina chlorotica (Ach.) Mull. Arg. (A) #61 and 59. * Porina chlorotica vat. persicina (Korb.) Zahlbr. (O) #50. This taxon is generally associated with calcareous rock. A short description of the variety is given by Duncan (1970). Verrucaria maura Wahlenb. ex Ach. (O) #62. Site 3. Sheltered inlet overhung by trees, near fish trap on trees or soil. Alectoria vancouverensis (Gyeln.) Gyeln. ex Brodo & D. Hawksw. (A) #79. This species is most easily distinguished from A. sarmentosa by its C+ red medullary reaction (due to olivetoric acid). There are morphological distinctions as well, all discussed by Brodo and Hawksworth (1977). Bryoria trichodes (Michx.) Brodo & D. Hawksw. subsp. americana (Mot.) Brodo & D. Hawksw. (Syn. Alectoria americana Mot.) #78 and 79 (A). The segregation of Bryoria from Alectoria is discussed in detail by Brodo and Hawksworth (1977). Cladonia chlorophaea (Florke ex Somm.) Spreng. (A) #71. Cladonia macilenta Hoffm. subsp. theiophila Asah. (C) #72. Cladonia scabriuscula (Del. ex Duby) Nyl. (O) #69. Cladonia subsquamosa (Nyl.) Vain. (A) #73. Cladonia transcendens (Vain.) Vain. (O) #74. THE CANADIAN FIELD-NATURALIST Vol. 91 Icmadophila ericetorum (L.) Zahlbr. (O) #70. Lepraria candelaris (L.) Fr. (O) #76. Peltigera horizontalis (Huds.) Baumg. (O) #68. Platismatia glauca (L.) W. Culb. & C. Club (R) #67. Sphaerophorus globosus (Huds.) Vain. (C) #66. Usnea longissima Ach. (A) #79. Site 4. Bark of Thuja plicata in coniferous forest. Calcium subquercinum Asah. (O) #84. This species is discussed by Tibell (1975) who reported it for North America for the first time. Chaenotheca brunneola (Ach.) Mull. Arg. (O) #85. Graphis elegans (Borr. ex Sm.) Ach. (O) #86. Icmadophila ericetorum (L.) Zahlbr. (O) #82. Lecanactis megaspora (Merr.) Brodo (A) #83. The taxonomy and nomenclature of this species is discussed by Brodo (1976). Spaerophorus melanocarpus (Sw.) DC. (R) #80. Site 5. On bark of Alnus rubra in forest openings. Arthonia radiata (Pers.) Ach. (O) #139. Arthothelium cfr. ilicinum (T. Tayl.) P. James (O) #87. The spores of this specimen were darker than is usual for the species. Arthothelium ilicinum was first reported for British Columbia by Brodo (1971) but apparently is not uncommon along the coast. Bacidia friesiana Korb. (O) #135. * Buellia griseovirens (Turn. et Borr.) Almb. (O) #117. Buellia griseovirens is often found sterile and is probably more common than existing collections would indicate. Both this specimen and a Swedish specimen with which it was compared contained norsticic acid (according to crystal tests with KOH + K,CO,) which is the basis for the KOH+ yellow-turning-red, PD+ yellow reactions of the soralia. The species is common in Great Britain on smooth-barked trees (Duncan 1970) and is probably a western American — western Europe disjunct like many other species (e.g., Letharia vulpina, Alectoria fremontii, Cladonia bellidiflora, etc.). Buellia penichra (Tuck.) Hasse (O) #116. Buellia punctata (Hoffm). Mass. (A) #94. Buellia stillingiana J. Stein (A) #138. Cetrelia cetrarioides (Del. ex Duby) W. Culb & C. Culb. (O) #107. Hypogymnia enteromorpha (Ach.) Nyl. (C) #93. Lecanora cfr. expallens Ach. (C) #134. The distribution of L. expallens was outlined by Brodo (1976) who reported it for the first time for British Columbia. This particular specimen, however, is aberrant in being esorediate and having broader spores; it may, in fact, represent a different species. The tallus does show the C+ orange reaction characteristic of L. expallens. Menegazzia terebrata (Hoffm.) Mass. (A) #91 and 122. Micarea melaena (Nyl.) Hedl. (O) #140. Nephroma laevigatum Ach. (A) #100 and 123. Parmelia sinuosa (Sm.) Ach. (O) #121. Parmelia sulcata Yayl. (A) #119. Parmeliella saubinettii (Mont.) Zahlbr. (C) #103. Peltigera collina (Ach.) Ach. (C) #106. Peltigera membranacea (Ach.) Nyl. (A) #99. 1977 Pertusaria ophthalmiza Ny\. (C) #92. This species is the one most frequently referred to by North American authors under the name P. multipuncta (Turn.) Nyl. Its taxonomy is discussed in detail by Dibben (1974). Pseudocyphellaria anomala Magn. (C) #115. Sticta limbata (Sm.) Ach. (O) #113. Thelotrema lepadinum (Ach.) Ach. (O) #136 and 97. Site 6. On bark of Acer macrophyllum. Caloplaca ferruginea (Huds.) Th. Fr. (O) #133. Lobaria oregana (Tuck.) Mull. Arg. (A) #131. Lobaria pulmonaria (L.) Hoffm. (A) #132. Ochrolechia oregonensis Magn. (A) #125. Pseudocyphellaria anthraspis (Ach.) Magn. (A) #130. Pseudocyphellaria aurata (Ach.) Vain. (O) #126. Pseudocyphellaria crocata (L.) Vain. (A) #127. Sticta fuliginosa (Dicks.) Ach. (O) #129. Sticta limbata (Sm.) Ach. (A) #124. Literature Cited Ahti, T. 1964. Macrolichens and their zonal distribution in boreal and arctic Ontario, Canada. Annales Botanici Fennici |: 1-35. Anonymous. 1971. Temperature and precipitation 1941- 1970, British Columbia. Atmospheric Environment Service, Department of the Environment, Ottawa. 94 pp. Bird, C.D. and R.D. Bird. 1973. Lichens of Saltspring Island, British Columbia. Syesis 6: 58-80. Brodo, I. M. 1971. Lichenes Canadenses exsiccati: A new series of Canadian lichens. Bryologist 74: 151-153. Brodo, I. M. 1976. Lichenes Canadenses exsiccati: Fas- cicle II. Bryologist 79: 385-405. Brodo, I. M. and D. L. Hawksworth. 1977. Alectoria and allied genera in North America. Opera Botanica 42: 1-161. Dibben, M. J. 1974. The chemosystematics of the lichen genus Pertusaria in North America north of Mexico. Ph.D. thesis, Duke University, Durham, North Carolina. 587 pp. Duncan, U. K. 1970. Introduction to British lichens. T. Buncle and Co., Arbroath, United Kingdom. 292 pp. NOTES 309 Halliday, W. E. D. 1937. A forest classification of Canada. Forest Service Bulletin 89. Canada Department of Mines and Resources, Ottawa. 59 pp. Harris, R.C. 1975. A taxonomic revision of the genus Arthopyrenia Massal. s. lat. (Ascomycetes) in North America. Ph.D. thesis, Michigan State University, East Lansing, Michigan. 288 pp. James, P. W. and A. Henssen. 1976. The morphological and taxonomic significance of cephalodia. Jn Liche- nology: Progress and problems. Edited by D. H. Brown, D. L. Hawksworth, and R. H. Bailey. Academic Press, London. pp. 27-77. Krajina, V. J. (Editor). 1965. Ecology of Western North America. Volume |. University of British Columbia, Vancouver, British Columbia. 112 pp. Otto, G. F.and T. Ahti. 1967. Lichens of British Columbia. Preliminary checklist. Processed publication, Department of Botany, University of British Columbia, Vancouver. 40 pp. Pike, L. H., W. D. Denison, D. M. Tracy, M. A. Sherwood, and F. M. Rhoades. 1975. Floristic survey of epiphytic lichens and bryophytes growing on old-growth conifers in western Oregon. Bryologist 73: 389-402. Poelt, J. 1969. Bestimmungeschlus sel europaischer Flech- ten. Cramer, Lehre. 757 pp. Rowe, J.S. 1959. Forest regions of Canada, Forestry Branch, Bulletin 123. Canada Department of Northern Affairs and Natural Resources, Ottawa. 71 pp. Thomson, J.W., G.W. Scotter, and T. Ahti. 1969. Lichens of the Great Slave Region, Northwest Territories, Canada. Bryologist 72: 137-177. Tibell, L. 1975. The Caliciales of boreal North America. Symbolae Botanicae Upsalienses 21: 1-128. Whitford, H. N. and R. D. Craig. 1918. Forests of British Columbia, Canada. Commission of Conservation, Ottawa. 409 pp. Received | June 1976 Accepted 5 March 1977 Narwhals (Monodon monoceros) Observed near King Christian Island, Northwest Territories NICHOLAS A. ROE and WILLIAM J. STEPHEN Beak Consultants Limited, 3530 — 11A Street N.E., Calgary, Alberta T2E 6M7 Roe, Nicholas A. and William J. Stephen. 1977. Narwhals (Monodon monoceros) observed near King Christian Island, Northwest Territories. Canadian Field-Naturalist 91(3): 309-310. The narwhal (Monodon monoceros) is commonly found in Canadian Eastern Arctic waters, but the peripheral parts of its summer range remain poorly documented (K. Hay, personal communication). Catch records indicate that it regularly occurs in Jones Sound, but the most westerly catch location is Resolute at 74°41’ N, 95°00’ W (Mansfield et al. 1975). On 6 September 1976, while flying at 61 m above ground level approximately 340 km northwest of Resolute, Stephen counted 10 narwhals ina patch 310 THE CANADIAN FIELD-NATURALIST of open water enclosed by ice approximately 0.4 kmin diameter at 77°20’ N, 103°30’ W in Maclean Strait between King Christian and Lougheed Islands, District of Franklin. Tusks were clearly visible on a number of the animals, thus confirming the identifica- tion. From published records it appears that narwhals penetrate further north into the Arctic than any other cetacean. In the Soviet Arctic, observations have been made between 81°00’ N and 84°40’ N (Rutilevskii 1958; Tomilin 1967), and schools in the European Arctic have been seen between 82°30’ N and 83°00’ N (Nansen 1897: Herbert 1969). In Canadian waters, narwhals probably spend the winter in open water in Baffin Bay, and move northward and westward as ice breaks up (Mansfield et al. 1975). Their summer range is centered at northern Baffin Island, but recent summer sight records have also been made around Grinnell Peninsula (Devon Island) and Penny Strait, where their numbers are not large (K. Hay, personal communication). Our record extends the summer range of the narwhal further westward and northward in the Canadian Arctic archipelago. The observation was made while the authors were Vol. 91 engaged in studies for Panarctic Oils Limited, Calgary, Alberta, whom we thank for permission to publish this note. We are also very grateful to K. Hay and D. E. Sergeant, Environment Canada, Fisheries and Marine Service, Arctic Biological Station, Sainte- Anne de Bellevue, Quebec, for information on Soviet and other records of narwhal distribution. Literature Cited Herbert, W. 1969. Across the top of the world. Longman, London. 209 pp. Mansfield, A. W., T. G. Smith, and B. Beck. 1975. The narwhal, Monodon monoceros, in eastern Canadian waters. Journal of the Fisheries Research Board of Canada 32: 1041-1046. Nansen, F. 1897. Farthest north. Archibald Constable and Company, Westminster, London. Volume 2. 671 pp. Rutilevyskii, G. L. 1958. A narwhal in the region of drifting station North Pole-5!. Problemy Arktiki, Sboriuk Statei 3: 116-119. (In Russian.) Tomilin, A. G. 1967. Mammals of the USSR and adjacent countries. Volume IX. Cetacea. Israel Program for Scientific Translations, Jerusalem. 756 pp. Received 7 February 1977 Accepted 16 July 1977 A Zone-tailed Hawk in Nova Scotia IAN A. MCLAREN! and ANDREW MACINNIS2 ‘Biology, Department, Dalhousie University, Halifax, Nova Scotia B3H 4J1 2Nova Scotia Department of Lands and Forests, Waverley, Nova Scotia BON 2S0 McLaren, Ian A. and Andrew MacInnis. 1977. A Zone-tailed Hawk in Nova Scotia. Canadian Field-Naturalist 91(3): 310-311. This report gives details of observations of an adult Zone-tailed Hawk (Buteo albonotatus) found in the vicinity of Musquodoboit Harbour, Halifax County, Nova Scotia, in autumn 1976. The bird was first seen on 24 September by MacInnis as it quartered along the estuary of the Musquodoboit River, flying over the water, riverside lawns, and clearings. Later in the day MacInnis saw it along a forested hillside. The day was dull and the bird appeared all black. In size and manner it gave MacInnis the impression of being harrier-like. On 25 and 26 September, during MacInnis’ absence from the vicinity, it was repeatedly seen near their house on the estuary by Eric Crowell and family, who reported that the bird had a distinctive white tail band. When MacInnis next saw the bird on 29 September he noted this tail band and other features that made him suspect it was a Zone- tailed Hawk. He then contacted McLaren who, along with others, saw the bird several times subsequently. We are thus able to add to our own observations from field notes kindly given to us by Roger Burrows, Ian MacGregor, Eric Mills, and Wayne Neily. All sightings were within an area extending about 1 km north, 2 km south, and 3 km east of the town center of Musquodoboit Harbour. The bird was seen, evidently actively searching at times, over shallow water, shores, scrubby fields, semi-cleared hillsides, lawns with and without shade trees, coniferous and mixed woods, and even over the commercial center of town. The bird was variously stated to be the size (or length) of a Marsh Hawk (Circus cyaneus), Red-tailed Hawk (Buteo jamaicensis), Rough-legged Hawk (Buteo lagopus), and larger than a Broad-winged Hawk (Buteo platypterus). Its flight was distinctive. During directed glides its wings were often held horizontally, usually bent at the carpal joint. But when it soared its wings were generally held above the horizontal, sometimes slightly flexed, but more often straight and harrier-like. Its flight when soaring was 1977 rocking and unsteady. When it soared the bird was indeed like a Turkey Vulture (Cathartes aura), as pointed out in some field guides; however, three observers familiar with the vulture thought the wing dihedral was not as great in the hawk. The bird was watched for several minutes on 2 October by McLaren as it worked along 2 km of wooded ridge, about 50-75 m at the summit. It flapped and glided along the ridge in both directions, periodically flapping hard to gain height and then pausing to soar in tight spirals 50-100 m above treetops. Twice it plunged rapidly from such a spiral to inspect or attack (?) something in the trees. The behavior seemed quite unlike that of any of the local raptors. Two attempts to capture prey were seen: Crowell saw the bird drop “like an Osprey” (Pandion haliaetus) into long grass near the estuary; and MacInnis saw it make a rapid feint at a small bird on the top of a roadside tree. Such swift attacks from a bird that soars like, and perhaps mimics, the Turkey Vulture have been commented upon by Willis (1966) and Zimmerman (1976). Although at a distance the bird appeared all black, many details were evident at ranges that at times were as small as 30 m. Some blackish parts were thought to have a brownish cast by Mills and MacGregor. The broad, grayish-white band across mid-tail was not as marked when the tail was closed, but was as conspicuous as depicted in field guides when the tail was slightly spread. The narrow, grayish-white basal tail bands (seen by five observers who reported two or “at least two”) were visible only when the bird was nearby. Only McLaren and Mills reported a dingy terminal tail band. The paler margins of the underwings, superficially like those of a Turkey Vulture, could not be seen ata distance in poor light. Among field guides for Central and North America, that of Peterson (1961), which shows a gray, not white or grayish-white, ground color of the primaries, seems to portray our bird accurately. Friedmann (1950) described the inner webs of the remiges as paling to deep neutral gray and to grayish white, barred with fuscous black. The fine barring was seen by three observers, and McLaren thought that barring extended to the black underwing coverts, not just part way down the remiges as implied in some field guides. The upper parts appeared to be uniformly dark to some, but Mills observed lighter areas at the base of the primaries and MacGregor and Neily saw the pale mid-tail band from above, perhaps when the tail was more open. The feet and legs were bright yellow according to all observers, and extended to at least the vent (Mills) or beyond the base of the tail (MacGregor). The legs were dropped briefly for three observers, and the long, NOTES 311 unfeathered tarsi had no bands or jesses (MacGregor). All observers were impressed with the large size and bright yellow color of the cere, seemingly larger than on a number of paintings of the bird consulted by us. On color slides there is a hint of whitish around the yellow on the face; possibly observers were failing to distinguish the cere from pale surrounding feathers. On 3 October, Crowell obtained a few color slides of the bird, using a 2x teleconverter with a 50-mm lens. The results seem to us to be convincing evidence for the occurrence of this distinctive bird (slides and prints to National Museum of Canada). The bird was seen next day briefly and for the last time by Neily. The species breeds in the United States only in Arizona, Texas, and New Mexico, and as far as we know strays have occurred only in California and Nevada. It is migratory in northern parts of its range. It can never be known if such an unprecedented bird was a true vagrant or an escaped captive, but the latter possibility can be examined further. Richard Ryan, an American expert on “escapes,” writes (personal communication) that he has “personally never seen nor heard of one in the zoo, pet or falconry trade” and that for birds in general “birds of the year constitute 80-90% of recently shipped captives.” An escape from the United States or elsewhere in Canada would involve subsequent disoriented flight to Nova Scotia (as for a stray), so that the possibility of escape in Nova Scotia is more likely. We know of no falconers in Nova Scotia, and the bird has not been kept in our “wildlife park” at Shubenacadie (L. Pace, personal communication). M. Sellars, Canada Department of Agriculture, informs us that no hawks have been imported via, or legally shipped through, Halifax International Airport in the months prior to our sighting. Although R. Ryan (personal communica- tion) notes that “birds are very, very rarely shipped by boat,” we have learned locally that birds are common as shipboard pets. These are checked and quarantined in Halifax, and we learn from E. D. Kay, Department of Agriculture, that inspectors have not seen hawks of any sort among them. Literature Cited Friedmann, H. 1950. The birds of North and Middle America. United States National Museum Bulletin 50. Part. 11. 793 pp. Peterson, R.T. 1961. A field guide to western birds. Houghten Mifflin, Boston. 366 pp. Willis, E.O. 1966. A prey capture by the Zone-tailed Hawk. Condor 68: 104-105. Zimmerman, D. A. 1976. Comments on feeding habits and vulture-mimicry in the Zone-tailed Hawk. Condor 78: 420-421. Received 24 December 1976 Accepted 5 May 1977 S12 THE CANADIAN FIELD-NATURALIST Summer Use of a Highway Crossing by Mountain Caribou DONALD R. JOHNSON! and MICHAEL C. TODD2 ‘Department of Biological Sciences, University of Idaho, Moscow, Idaho, USA 83843 2College of Forestry, Wildlife and Range Sciences, University of Idaho, Moscow, Idaho, USA 83843 Johnson, Donald R.and Michael C. Todd. 1977. Summer use of a highway crossing by mountain caribou. Canadian Field-Naturalist 91(3): 312-314. Abstract. Caribou use of a highway crossing point near Kootenay Pass, British Columbia was monitored with a time-lapse camera during the summer months when highway traffic was heaviest. Caribou approached the crossing on at least 11 occasions throughout the daylight hours, including times of peak traffic flow. The number of approaches declined as the season progressed. Additional approaches undoubtedly occurred during the daylight hours, but these were not recorded by the camera, and during periods of darkness when the camera was inoperative. We conclude that mountain caribou have become habituated to the presence of the highway and road traffic and that they continue to use a traditional movement route despite man-caused harassment and mortality. Although caribou reaction to newly-placed move- ment barriers has received recent attention (Miller et al. 1972; Child 1973), little information is available on the reaction of caribou to permanent, man-made obstacles such as a heavily used highway. Klein(1971) briefly mentioned that wild reindeer terminated use of a range area a few years after construction of a main highway and railroad in Norway. Bergerud (1974) has discussed the reaction of caribou to visual, auditory, and olfactory stimuli including man-caused distur- bances. Mountain caribou, Rangifer tarandus montanus, have moved across heavily-traveled British Columbia Highway 3 near Kootenay Pass since the highway was completed in 1963. Most caribou have been observed crossing the highway at three locations both east and west of the summit (Freddy 1974, p. 40). Considering topographic features, it is likely that these crossings represent sites at which the highway intercepts traditional movement routes. Methods We monitored caribou approaches to one of these crossing points (North Fork of Summit Creek, 4 km east of Kootenay Pass) from 18 June through 23 August 1976 using a time-lapse movie camera in order to determine (1) the number and composition of caribou using the crossing, (2) the time of day the approaches occurred, (3) the pattern and frequency of approaches in relation to traffic flow, and (4) the reaction of caribou to the presence of motorists onthe highway. A Minolta D-6 Super 8-mm movie camera housed in a protective cover was mounted ona tree 5 mabove the ground so that it monitored about 0.5 km of highway. The camera was powered by a 6-V rechargeable battery and fitted with instrumentation to regulate automatically exposure rate (Four Seasons Services, Laramie, Wyoming). A light sensor limited the camera operation to daylight hours. Through use of an exposure rate of | frame/ minute, 800-1000 frames were exposed daily, depending on day length and cloud cover at dusk and dawn. At this exposure rate, a 3600-frame cartridge of color film (50 ft) was expended in 3.6 to 4.5 days. Because of logistical problems, we chose to monitor the site at weekly intervals rather than continuously. But monitoring occurred during some weekends when traffic was heaviest as well as during weekdays. Exposed film was examined in a viewer and the presence of caribou and traffic on each frame was recorded. Slow-moving or stopped vehicles were recorded more than once since they represented a continuous disturbance at the site. The time of each approach and its duration were calculated from the time exposure commenced and the exposure rate. Results and Discussion During more than 570 h of camera operation, the occurrence of caribou at the crossing was confirmed on I! occasions (Table 1). In 10 instances these represented either one or two animals; some of these could be identified as bulls, based on body and antler size. The cow-calf band, which has remained in the Summit or Carolina Creek basins during recent summers, was photographed at the crossing on 30 July. Because of the possibility of repeat approaches by the same individuals, it is difficult to estimate the total number of caribou photographed at the site. Based on body size and the presence or absence of antlers, a minimum of 12 different animals was photographed (two large bulls, seven cows and subadults, three calves). Since the total number of caribou in the West Kootenay band is estimated at 25-30 animals (Johnson 1976), perhaps one half of these were photographed at this traditional crossing during the monitoring period. Caribou approaches to the crossing occurred throughout the daylight hours (0618-1940 hours), NO Fa TABLE 1—Time of day, number and composition of caribou photographed on Highway 3. Only if animals were in certain positions could we see and record whether they were antlered or antlerless Date Time Number and composition 19 June 1140-1142 1 antlered bull 20 June 1158-1621 2 antlered bulls 1808-1940 2 subadults 25 June 1545 2 subadults 1746 1 subadult 26 June 0618 1 subadult 0657-0658 1 subadult 12 July 1248-1252 2 subadults 25 July 1052 1 antlerless cow; | calf 30 July 1007-1029 7 adults and subadults; 3 calves 21 August 1806 2 adults, anterless including times of peak traffic flow (Figure 1). Caribou usually appeared only briefly within camera view. On 20 June, however, four caribou appeared in 27 frames over a 2-h period (Table 1). On several occasions caribou were photographed while they were licking at the surface of the highway, perhaps at oil spots. Loggers have reported that caribou licked grease fittings on their machinery parked overnight (Layser 1974). There was a progressive increase in_ traffic throughout the summer. If we ignore days when stalled vehicles inflated the total, 29% of the frames FRAMES WITH MOTOR VEHICLES 0800 1000 1200 TIME NOTES SS exposed in June contained at least one motor vehicle; 36% of those exposed in July and 39% of those exposed in August contained at least one vehicle. There was a progressive decrease in the number of frames containing caribou during the same period. In June, caribou were found in 34 frames during four days: in July they occurred in eight frames during three days; and in August their presence was detected in only a single frame. We believe that this decrease in the use of the crossing by caribou was not related to increased traffic flow since caribou were not deterred from the crossing at times of peak traffic flow (Figure 1). It is more likely that caribou moved to higher elevations as the summer progressed and ground forage became available after snow melt. Motorists frequently stopped at the crossing because of car trouble, to pick berries, or to view caribou. On several occasions the camera recorded vehicles stopped along the highway and a number of people standing on the shoulder of the road. Caribou were often photographed soon after the motorists moved on. The presence of several stopped vehicles indicated that caribou probably were present along the highway on 27 July and 10 August but no animals were photographed on these days. The limitations of a single camera with a l-min exposure rate are obvious. We have also confirmed the use of the crossing at night although we do not know if the frequency of use differs between daylight and darkness. Our direct observation of caribou behavior at the crossing during the summer months indicated that 'g00 1400 1600 2000 OF DAY FIGURE |. Caribou approaches (singly or in groups) to the North Fork Highway Crossing in relation to traffic flow. Numbers indicate approaches within time intervals. 314 THE CANADIAN FIELD-NATURALIST some animals approach the highway with caution and seek cover in the nearby timber if motorists stop to view them. Others remain alert but continue to feed beside the highway or move off slowly in response to the presence of motorists nearby. At least seven caribou have been killed in collisions with motor vehicles since the highway was opened, including a cow and calf in separate incidences at the North Fork Crossing in 1976. Caribou have been shot illegally along the highway (Freddy 1974). Despite this harassment and mortality they continue to cross the highway at what appears to be traditional locations. Completion of British Columbia Highway 3 through the Kootenay Pass region has fostered the development of a utility corridor along its route including a natural gas pipeline and two power lines (Johnson 1976). Although restriction of these devel- opments to a narrow corridor localizes their environ- metal impact, the continued increase in the number of these hazards may eventually interrupt the normal north-south movement of caribou in this region. The impact of these developments on caribou movement needs further study. We thank the USDA Forest Service, the Wash- ington Game Department, and the West Kootenay Outdoorsmen for their continued support during this study. Jeff Yeo and Michael D. Johnson provided field assistance. Vol. 91 Literature Cited Bergerud, A. T. 1974. The role of the environment in the aggregation, movement and disturbance behavior of caribou. Jn The behaviour of ungulates and its relation to management. Edited by V. Geist and F. Walther. IUCN Publication, New Series 24. 2 volumes. pp. 552- 584. Child, K. N. 1973. The reactions of barren-ground caribou (Rangifer tarandus granti) to simulated pipeline and pipeline crossing structures at Prudhoe Bay, Alaska. Alaska Cooperative Wildlife Research Unit Completion Report. 49 pp. Freddy, D. J. 1974. Status and management of the Selkirk caribou herd, 1973. M.Sc. thesis, University of Idaho. 132 pp. ; Johnson, D. R. 1976. Mountain caribou: threats to sur- vival in the Kootenay Pass Region, British Columbia. Northwest Science 50: 97-101. Klein, D. R. 1971. Reaction of reindeer to obstructions and disturbances. Science (Washington) 173: 393-398. Layser, E. F., Jr. 1974. A review of the mountain caribou of northeastern Washington and adjacent northern Idaho. Journal of the Idaho Academy of Science, Special Research Issue 3. 63 pp. Miller, F. L., C. J. Jonkel, and G. D. Tessier. 1972. Group cohesion and leadership response by barren-ground caribou to man-made barriers. Arctic 25: 193-202. Received 18 January 1977 Accepted 15 June 1977 Changes in the Avifauna of the West Foxe Islands, Northwest Territories, 1956-1976 F. G. COOCH Canadian Wildlife Service, Ottawa, Ontario KIA 0M8 F. G. Cooch. 1977. Changes in the avifauna of the West Foxe Islands, Northwest Territories, 1956-1976. Canadian Field-Naturalist 91(3): 314-317. Abstract. During July 1976, a resurvey was made of the bird populations of the West Foxe Islands near Cape Dorset, Northwest Territories. Three species new to the area were recorded: Great Black-backed Gull, Eastern Kingbird, Yellow- rumped Warbler. Changes in numbers and status of other species since the last survey in 1956 were noted. Heavy snow during the winter of 1975-76 and delayed melt are considered to be causative factors in the changes in numbers of most species. Macpherson and McLaren (1959) published an annotated list of the birds of the southern Foxe Peninsula, Baffin Island (64° 14’ N, 76°13’ W), based on their observations of 1954 and 1955 and those of Cooch in 1955 and 1956 (Cooch, F. G. 1957. Birds observed in the vicinity of Cape Dorset, Baffin Island, in the summers of 1955 and 1956. Typed report, on file National Museum of Canada. 13 pp.). In the period 8-19 July 1976, while on a resurvey of the Northern Eider (Somateria mollissima borealis) populations of the Cape Dorset Migratory Bird Sanctuary, I recorded a number of species new to Cape Dorset, as well as changes in population status of species noted previously. The West Foxe Islands (Figure 1), South Island in Andrew Gordon Bay, and Sakkiak Island near Cape Dorset, were set aside asa 1977 HONTING ‘@: BILDFELL oo a NESFIELD x CUO MUNUSSON ; ‘9 MG, INNUKSHUK i Des 7 RUSSELL SCHIOLER : COATESWORTH =~ LUKE ~ <— CAPE DORSET NOTES 315 ALAREAK FIGURE |. West Foxe Islands, Northwest Territories (64°14’ N, 76°13’ W). Federal Migratory Bird Sanctuary in 1957 to provide protection for eiders. Spring breakup in 1976 in southern Baffin Island was atypical. May was abnormally warm and the above-normal snowfall began to melt rapidly. The weather changed dramatically in June when high winds, fog, and freezing rain were normal. One result of the high winds was the disappearance of much of the land-fast sea ice before mid-June. The unusual nature of the spring phenology, the heavy snow cover which persisted in many areas even after 20 July 1976, and the persistence of ice on most inland lakes apparently had a marked effect on the subsequent distribution and migration patterns of birds nesting in the Cape Dorset area and farther north. The following is a list of birds observed on the West Foxe Islands, 8-19 July 1976, with general comments from the Cape Dorset Area. COMMON LOON (Gavia immer) Uncommon spring transient 1955; relatively common summer resident 1976. Not known to breed on small tarns on coastal islands. Large inland lakes normally used by this species were still frozen on 18 July 1976. The continued presence of ice on the larger inland lakes must have affected the distribution of all three species of loons normally breeding in the Cape Dorest area. ARCTIC LOON (Gavia arctica) Uncommon spring transient in 1955 and 1956: relatively abundant — at least 15 individuals were recorded daily in July 1976 among the West Foxe Islands, where they had not been detected in summer in 1955 or 1956. RED-THROATED LOON (Gavia stellata) Common spring transient in 1955 and 1956; abundant in 1976 in the West Foxe Islands. One nest was located ona I- ha tarn on Innukshuk Island. Previously not known to breed. CANADA GOOSE (Branta canadensis interior) (Branta canadensis hutchinsii) The West Foxe peninsula supports two races of Canada Geese, medium-sized form (B.c. interior) and a small form (B.c. hutchinsii). Usually the latter breeds along the north coast of the Foxe Peninsula from Harkin Bay northeast along the Great Plains of the Koukdjuak to Taverner Bay (67° 12’ N, 72°25’ W) whereas the larger race is restricted to the coastal area south of Cape Queen. In 1976 both races nested in the West Foxe Islands where neither had been recorded breeding previously. On 10 July 1976, an intermediate-sized pair with two young approximately 3 days old were found on the top of Luke Island, and on 14 July a pair representing the small race was observed with three young on Blades Island. This is taken as further evidence of displaced nesting in 1976. ATLANTIC BRANT (Branta bernicla hrota) Cooch (manuscript) recorded a successful nesting attempt by the species on the West Foxe Islands in 1956. At least four pairs attempted to nest there in 1976 and others were reported by Inuit to be nesting at widely scattered locations along the south coast of Baffin Island. The nearest known Brant nesting colony is at Cape Dominion 300 km north of the West Foxe Islands. An apparent change in autumn distribution was reported by Inuit who camp at the head of Andrew Gordon Bay about 80 km east of Cape Dorset. Several thousand Brant are nowseen regularly in late August feeding in the sea and along the tidal wrack at the mouth of the Saunders River. This autumnal build-up did not occur in 1955 and 1956 and apparently started only in the late 1960s. 316 LESSER SNOW GOOSE (Anser caerulescens) Although an abundant spring and autumn migrant through the area along a broad front from Cape Queen east to Markham Bay, it does not normally breed locally. A nest containing parts of three eggs was detected on Dune Island 12 July 1976, and on 14 July 1976 a mixed pair (white phase male-blue phase female) accompanied by a single white phase gosling was seen on nearby Innukshuk Island. Inuit reported this species nesting in non-colonial situations at various places along the south coast in 1976. They had previously noted much displaced breeding in 1972, when the colony area at Bowman Bay was snow-covered into early July, as it was in 1976. They also noted that the peculiar migration pattern of 1972 was repeated at Cape Dorset in 1976. Normally the geese make a rapid straight-line flight from Ungava to the vicinity of Cape Dorset — Andrew Gordon Bay, then turn northeast toward Bowman Bay and the Great Plains of the Koukdjuak. In both 1972 and 1976 significant reverse migrations were observed a few hours after the first flights passed over Cape Dorset. Subsequent flights reaching Cape Dorset appeared confused and veered to the northwest, toward the flat coastal tundra near Cape Dorchester, which was apparently relatively snow-free. PINTAIL (Anas acuta) Three males were seen on a small lake on Blades Island 13 July 1976. The species is not normally found in the Cape Dorset area. Northward post-breeding dispersal of males of the species is a widespread phenomenon. NORTHERN EIDER (Somateria mollissima borealis) After the Black Guillemot, this is the most abundant species nesting in the West Foxe Islands. Estimates of breeding pairs on the West Foxe Islands were 982 in 1955, 1295 in 1956, and in 1976 only 367 pairs. Almost all of the decrease occurred on Tunitjuak Island where fewer than 50 pairs were found in 1976 compared to 667 in 1955 and 755in 1956. Although much of this decrease must be attributed to depredations by Inuit taking eggs and shooting female eiders (Cooch, unpublished data), the heavy snow cover of the winter of 1975-76 and the slow thawing of interior lakes were also undoubtedly important factors contributing to the decline of the numbers of eiders on the largest of the West Foxe Islands. Egg-laying effectively ceased on 20 July 1976 (P. Putagook, personal communication), about 4 days later than the previously recorded last date of 16 July 1956. Predation was unusually heavy because of the numbers of non-breeding Herring Gulls, Glaucous Gulls, and Parasitic Jaegers. OLDSQUAW (Clangula hyemalis) An abundant spring migrant but normally an uncommon summer resident. Several flocks of up to 25 birds were seen daily in July 1976. At least three pairs of Oldsquaws nested on the inner West Foxe Islands in 1976; none had been recorded breeding in 1955 or 1956. SEMIPALMATED PLOVER (Charadrius semipalmatus) This most abundant species of shorebird breeding in the sanctuary 1s found primarily in dry upland lichen and grassy swales. First hatching detected in 1976 was on 16 July on Coatesworth Island. Population numbers were little changed from previous surveys in 1955 and 1956. THE CANADIAN FIELD-NATURALIST Vol. 91 PURPLE SANDPIPER (Calidris maritima) An abundant spring and autumn migrant and an uncommon summer resident. Not previously known to breed on the West Foxe Islands. Abundant summer resident in 1976; breeding was strongly suspected on the basis of distraction display by three widely scattered pairs. At least 250 individuals were resident in July 1976 compared to none in July 1955 or 1956. SEMIPALMATED SANDPIPER (Calidris pusilla) Abundant around tarns in 1955 and 1956 and a common breeding bird. A single individual was seen in 1976, with no evidence of breeding. PARASITIC JAEGER (Stercorarius parasiticus) Parasitic Jaegers are not known to breed in the West Foxe Islands. In 1955 only a single bird was seen. In 1956, when there was a relatively late spring, more than 150 sightings were made during the period 15 May — 10 September. In 1976 in excess of 200 sightings were made in the period 9 to 16 July alone. The presence of Parasitic Jaegers in numbers on the West Foxe Islands in 1976 is probably related to the non- breeding of Lesser Snow Geese on the Foxe Basin Coast of Baffin Island. A similar increase in numbers was observed in 1956 when snow covered the Great Plains of the Koukdjuak and Southampton Island until 22 June. In years of long- lingering snow, non-breeding of geese, lows in the lemming populations, and lack of suitable nesting habitat, jaegers disperse widely in search of alternative feeding sites. GREAT BLACK-BACKED GULL (Larus marinus) A single representative of this species was seen at Ooglukjuak Island 14 July 1976. It has been seen increasingly on the Atlantic coast of Baffin Island in recent years (D. N. Nettleship, personal communication), but has not been reported so far west on the south coast. HERRING GULL (Larus argentatus smithsonianus) In 1955 and 1956, 15 scattered pairs nested on the West Foxe Islands. No nesting was detected in July 1976, although 15—20 birds were seen over the islands at any given time. A communal roost on the east end of Tunitjuak Island is further evidence of non-breeding in 1976. The almost total lack of gulls of any species in the vicinity of Cape Dorset harbor was startling when compared to their abundance in 1955 and 1956. KUMLIEN’S GULL (Larus glaucoides kumlieni) The colony at Ooglukjuak (Macpherson and McLaren 1959; Macpherson 1961) numbered 28 breeding pairs on 14 July 1976, plus 24 potential pairs, a net decrease of 50 pairs since the last census on 18 July 1956 (Cooch, manuscript). Of 36 eggs candled, none gave an indication of hatching before 21 July. THICK-BILLED MURRE (Uria lomvia) In the period 1955-56 Brunnich’s Murres, presumably from the vast colony at Digges Island (120 kmsouth of Cape Dorset) were noted in the vicinity of Cape Dorset in a restricted area between Dorset Island and Neta Island 16+ km (10 mi) to the east. They were most abundant in May and early June along the edge of the land-fast ice. In 1976, murres, although most common in their traditional area, were seen commonly at least as far east as South Island, Andrew Gordon Bay, approximately 128 km (80 mi) east of 1977 Cape Dorset. This may reflect a poor breeding season at Digges Island or, more probably, the abundance of non- breeding yearlings produced in the excellent 1975 breeding season. Unusually extensive areas of open water occurring throughout the Neta, Shemia, and West Foxe Island chains before 15 June may also have attracted unusually large numbers of murres. At least 4000 birds were in the vicinity of the West Foxe Islands between 9 and 19 July 1976. BLACK GUILLEMOT (Cepphus grylle ultimus) An abundant nesting bird wherever suitable talus or scree slopes are available. The major colony site on Russell Island remained almost completely snow-covered in 1976 until my departure from the islands on 17 July. No young were observed and many clutches contained a single egg or reached their full complement of two during the interval 14-16 July. The peak of hatching was not expected until the second week of August. The long-persisting snow banks which covered many choice talus slopes radically re- distributed guillemots throughout the islands, forcing some into inland crevices overlooking freshwater ponds — sites which were not utilized in 1955 or 1956. No overall decrease in the total of guillemots was detected in the West Foxe Islands although the colony on Russell Island was reduced to perhaps 500 pairs, down from the 3000 recorded in 1956. EASTERN KINGBIRD (7 yrannus tyrannus) A new record for the Cape Dorset area and apparently for Baffin Island. A solitary male was seen feeding on masses of newly hatching mosquitoes on Blades Island 13 July 1976. WHEATEAR (Oenanthe oenanthe) A nest containing three eggs was discovered ina crevice on a high rocky outcrop on Tunitjuak Island on 14 July 1976. Wheatears are not uncommon on the south coast of Baffin Island, approximately 80km (50 mi) south of Lake Harbour. Macpherson and McLaren (1959) reported three breeding pairs of Wheatears nest-building at the Aitken Lakes near the Cape Dorset settlement. NOTES 317 YELLOW-RUMPED WARBLER ( Dendroica coronata) A new record for the Cape Dorset area and apparently for Baffin Island. A solitary male was seen feeding on mosquitoes not far from the site where the Eastern Kingbird was seen earlier on 13 July 1976. WATER PIPIT (Anthus spinoletta) A common breeding species in both 1955 and 1956, totally absent in 1976. COMMON REDPOLL (Acanthis flammea) Common spring migrant, relatively uncommon summer resident in 1955 and 1956. None seen in 1976. LAPLAND LONGSPUR (Calcarius lapponicus) An abundant breeding species in 1955 and 1956 but virtually absent in 1976 when only one pair and two stray males were seen. SNOW BUNTING ( Plectrophenax nivalis) In 1955 and 1956 Snow Buntings nested abundantly on every island in the West Foxe chain, at densities approaching one pair per hectare. In 1976 this crevice-nesting species was severely affected by long-lingering snow drifts in favored nesting sites and densities dropped to one pair per 10 hectares. A similar lack of Snow Buntings was recorded in the vicinity of Cape Dorset settlement. Literature Cited Macpherson, A.H. 1961. Observations on Canadian Arctic Larus gulls, and on the taxonomy of L. thayeri Brooks. Arctic Institute of North America Technical Paper Number 7. 40 pp. Macpherson, A. H.and I. A. McLaren. 1959. Notes on the birds of southern Foxe Peninsula, Baffin Island, North- west Territories. Canadian Field-Naturalist 73(2): 63-81. Received 20 December 1976 Accepted 16 July 1977 Unusual Predators of Snow Goose Eggs KENNETH F. ABRAHAM, PIERRE MINEAU, and FRED COOKE Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6 Abraham, Kenneth F., Pierre Mineau, and Fred Cooke. 1977. Unusual predators of Snow Goose eggs. Canadian Field- Naturalist 91(3): 317-318. Several bird and mammal species prey on the eggs of Lesser Snow Geese (Anser caerulescens caeru- lescens). Parasitic Jaegers (Stercorarius parasiticus), gulls (Larus spp.), and arctic fox (Alopex lagopus) are considered the most important predators (Cooch 1958; Ryder 1969a). Less important predators include Sandhill Cranes (Grus canadensis) (Harvey et al. 1968), wolves (Canis lupus), polar and grizzly bears (Ursus maritimus and U. horribilis) (F. G. Cooch, Canadian Wildlife Service, personal communication; Barry 1967). In this note we describe predation on Snow Goose eggs at La Pérouse Bay, Manitoba, by two previously unreported species. La Pérouse Bay is located 40 km east of Churchill, Manitoba. Caribou (Rangifer tarandus) occur near many Snow Goose colonies and are seen regularly at La Pérouse Bay, but have never been reported as goose- egg predators. On 8 June 1976, KFA observed a young caribou walking from island to island in a shallow lagoon where Snow Geese and Common 318 Eiders (Somateria mollissima) nest. Female geese and eiders flushed from nests as the caribou moved within 10 min of the caribou’s leaving. For 30 min the female The caribou ate willow (Salix spp.) buds and leaves on several islands, but on one it stopped by an active Snow Goose nest that we had marked, pawed at the nest, and appeared to eat something. The nest was checked immediately and five of six eggs were found crushed with little of their contents remaining. During the 5 min the caribou remained at the nest, the attendants could not be identified from among a group of 45 yearlings and adults which gathered near the island, but a pair of Snow Geese returned within 10 min of the caribou’s leaving. For 30 min the female poked at the egg remains while the male stood 2 m away. Ihe female was observed to lower, then raise and tip back her head as if drinking, and also to eat some of the eggshells. (Ryder (1969b) discusses the significance of egg-eating by Snow Geese.) The nest was revisited on 10 June and had been partially reconstructed with the intact egg in the bowl. On 15 June the nest was empty; the fate of the last egg is unknown. On 15 June, KFA observed another young caribou similarly disturb two occupied nests about 3 km away from the location of the first incident, but in a much drier nesting area. Again the geese flushed ahead of the caribou and did not attempt to defend their nests. At the first nest, this caribou rolled the eggs out of the nest bow] with its right foreleg, crushed one and licked the egg, but did not eat the 15- to 20-day-old embryo. It then immediately moved to another nest 50 m away, repeated the sequence but this time picked up and ate one embryo. It continued to walk through the area, but was not seen at any more nests. A nest check showed one crushed and four intact eggs at the first nest, three crushed and two intact eggs at the second nest. Size and antler characteristics assured us that two different caribou were responsible for these acts of predation. Hunger may have been a motivating factor, but does not by itself explain it as in each case only part of the egg was eaten. As both appeared to be young individuals, perhaps curiosity was partially responsible for this behavior. Kelsall (1968) reviews the miscellaneous foods of reindeer and caribou, which include birds’ eggs and possibly nestlings, but stresses the rarity of these events. The reaction of the geese also indicates that caribou weren’t recognized as predators, although PM observed geese threatening and charging caribou during late incubation in 1975. We think it unlikely that caribou predation of Snow Goose eggs is of major importance. To our knowledge black bears (U. americanus) have never been sighted at a Snow Goose colony. Two THE CANADIAN FIELD-NATURALIST Vol. 91 sightings were made on 22 and 23 June 1975 at La Pérouse Bay, probably involving the same individual. Tracks indicated that several nests had been visited, but all the eggs had already hatched. On 23 June, PM witnessed one incidence of predation. The bear walked to an occupied nest with three eggs which had been incubated at least 16 days. As the bear neared the nest, the gander briefly adopted a high-intensity threat posture with wings extended and head held low before retreating with the female to a shallow stream 15 m away. The bear spent less than a minute at the nest. A subsequent check revealed that two of the three eggs had been eaten, the embryos having been removed from the crushed shells. The third egg was rotten; this may explain why it was not eaten. The goose continued to incubate for 2 days at which time we removed the egg from the nest. The omnivorous habits of black bears are well documented and they are known to supplement their diet with birds’ eggs. Given the present southward expansion of the Snow Goose breeding range (Dzubin et al. Blue and Snow Goose distribution in the Mississippi and Central flyways, Volume |, Canadian Wildlife Service Report, 1973), and the increasing tendency of black bears to move north of the tree line (Jonkel and Miller 1970), it is possible they will become more frequent predators at the more southern Snow Goose colonies. A continuing study of Snow Geese at La Pérouse Bay, Manitoba, is funded by the Canadian Wildlife Service, National Research Council, the Canadian National Sportsmen’s Show, the Wildlife Manage- ment Institute, and the Manitoba Department of Mines, Energy and Environmental Management. Literature Cited Barry, T. W. 1967. The geese of the Anderson River delta, Northwest Territories. Ph.D. thesis, University of Alberta, Edmonton. 212 pp. Cooch, F. G. 1958. The breeding biology and management of the blue goose (Chen caerulescens). Ph.D. thesis, Cornell University, Ithaca, New York. 235 pp. Harvey, J.M., B.C. Lieff, C.D. MacInnes, and J.P. Prevett. 1968. Observations on behaviour of sandhill cranes. Wilson Bulletin 80(4): 421-425. Jonkel, C.J. and F.L. Miller. 1970. Recent records of black bears (Ursus americanus) on the barren grounds of Canada. Journal of Mammalogy 51(4): 826-828. Kelsall, J. P. 1968. The migratory barren-ground caribou of Canada. Canadian Wildlife Service Monograph. 355 pp. Ryder, J. P. 1969a. Nesting colonies of Ross’s Goose. Auk 86(2): 282-292. Ryder, J. P. 1969b. Egg-eating by wild Lesser Snow Geese. Avicultural Magazine 75(1): 23-24. Received 19 November 1976 Accepted 12 April 1977 Sy NOTES Sy/19) New and Notable Finds in the Alaskan Vascular Flora G. HALLIDAY Department of Biological Sciences, The University, Lancaster LAI 4YQ, England Halliday, G. 1977. New and notable finds in the Alaskan vascular flora. Canadian Field-Naturalist 91(3): 000-000. Abstract. Plewropogon sabinei R. Br., Potentilla rubricaulis Lehm., and Pedicularis hirsuta L. are reported for the first time from Alaska; second records are given for Puccinellia angustata(R. Br.) Rand. & Redf. and Colpodium vahlianum (Liebm.) Nevski together with a number of important extensions of range. Twelve species are listed which have not previously been recorded from Prudhoe Bay, on the north-east coast. Key words: Pleuropogon sabinei, Potentilla rubricaulis, Pedicularis hirsuta, Alaskan flora The more interesting finds of vascular plants made during a brief visit to Alaska in July 1975 are listed below. Most of the collections were from the following localities. 1. Fairbanks — vicinity of University campus and the airfield (64°50’ N, 147°45’ W). 2. Livengood (80 km north-west of Fairbanks). (a) East-west ridge north of the Elliott High- way and 55 km south-west of Livengood (65°15’ N, 149°30’ W), (b) Brown Lake and nearby West Fork of Tolovana River, 13 km south-west of Livengood (65° 27’ N, 148°40’ W), (c) Livengood Dome (65°36’ N, 148°30’ W). 3. Wickersham Dome (between Fairbanks and Livengood) (65° 13’ N, 148°05’ W). 4. Sourdough Creek (80 km north-east of Fair- banks), north of the Steese Highway, and from head of creek to Point 5043’ (65°20’N, 146°30’ W). 5. Prudhoe Bay — between Kuparuk River and Sagavanirktok River (70°20’ N, 148°30’ W). 6. Galbraith Lake (north side of Brooks Range on trans-Alaska pipeline) — limestone hills east of the lake (69°30’ N, 149°30’ W). Of these areas one of the most interesting was 2(a). Several montane species were found on this ridge which were not previously known to occur so far west in the mountains between the Yukon and Tanana Rivers. Immediately north of the ridge lies the remote and botanically unknown area around the Wolverine and Sawtooth Mountains, an area which would certainly repay study. The species are listed as far as possible according to the sequence and nomenclature adopted by Hultén (1968). Herbaria are referred to by their international abbreviations. A set of nearly all the 500 numbers collected is in the Arctic Herbarium at LANC. Pleuropogon sabinei R. Br. Prudhoe Bay: north side of the Spine Road just west of Gathering Center-1. 12 July 1975, no. A223/75, ALA, BM, E, LANC, S. It was growing in bare mud by a small stream only a few yards from the road and associated with Arctophila fulva, Dupontia fisheri, and Eriophorum angustifolium. The population covered only a few square metres and the site had almost certainly been affected to some extent by the construction of the road. The plants were flowering freely. This is the first recard of this species from Alaska. Although the species was mapped by Hultén (1958) as an amphi-Atlantic species, he expressed the Opinion that recent finds on Wrangel Island and extreme eastern Siberia, and on Banks Island and Victoria Island, made it a circumpolar species. The Alaskan locality perfectly bridges the remaining gap between these eastern and western limits and vindi- cates Hultén’s (1968) faith in including this species in his Alaskan Flora. Puccinellia angustata (R. Br.) Rand. & Redf. Prudhoe Bay: pingo at south-east end of British Petroleum Operations Center reservoir. 9 July 1975, no. Al83/75, ALA, CAN, LANC, S. Along the western shore of the Sagavanirktok River estuary and on the adjacent tundra. I! July 1975, no. A213/75, LANC, S. At the latter locality P. angustata was a con- spicuous species along the beach and on level, clay polygon tundra nearby, where it was associated with Potentilla pulchella. The plants were prostrate and readily identified by the ciliolate margins and hairy lower half of the lemmas. This circumpolar species has a very dissected distribution in eastern Siberia and north-west North America. Tzvelev (1964) gives its eastern limit in Siberia as the New Siberian Islands and he specifically says it is absent from Wrangel Island and the Chukotsky Peninsula. Hultén (1968) shows it as occurring in the two latter areas, and in LANC there is an unnamed specimen distributed by LE and collected from Wrangel Island in 1967 by Petrovsky. With its small glumes and anthers, pilose lemmas and scabrous panicles it is clearly P. angustata. Previously the 320 westernmost locality in North America was thought to be Barter Island. Colpodium vahlianum (Liebm.) Nevski Prudhoe Bay: west side of the Sagavanirktok River estuary. 9 July 1975, no. A214/75, ALA, LANC. Only a few plants were found. They were growing in bare mud by a brackish inlet and were conspicuous by their bright green color, compared with the glaucous color of species of Puccinellia. The white, undulate roots were also characteristic. According to Tzvelev (1964) this species has its eastern limit in Siberia on the Taymyr peninsula and material from Wrangel Island he refers to the new species Puccinellia colpodioides Tzvelev, known only from this island. Unfortunately, the Wrangel Island locality appears in Hultén (1968). Puccinellia col- podioides is usually easily recognized by its long anthers, 1.5-2.5 mm, compared with 0.8-1.3 mm in C. vahlianum. The difference in glume length (and consequently the ratio between glume and lemma length) which Tzvelev (1964) cites, is erroneous. For P. colpodioides he gives the lengths of the outer and inner glumes as 1.8 mm and 2.5 mm respectively, but two recent collections of this species in LANC and distributed by LE had glume lengths of 2.1-2.7 mm and 2.8-3.4mm respectively. These ranges are indistinguishable from those of C. vahlianum. Like Pleuropogon sabinei, C. vahlianum has previously not been reported west of Banks Island. But it has recently been discovered in Alaska in the Brooks Range (D. F. Murray 1975, personal com- munication). Carex peckii Howe Fairbanks: campus road by forest margin west of the University. | July 1975, no. A20/75, CAN, LANC. Livengood: north end of Brown Lake, 13 km south- west of Livengood. 3 July 1975, no. A49/75, ALA, C, E, LANC, S. At the Livengood locality the sedge was growing in a grassy clearing by a track and near the lakeshore. Hultén’s (1968) map of this North American sedge shows a single locality in Alaska (Fairbanks) and one in Yukon (Dawson), but it has recently been reported by Scotter and Cody (1974) from the Mackenzie Mountains. Minuartia biflora (L.) Schinz. & Thell. Sourdough Creek: in snow-patch gully at head of creek, 980 m (3000’). 15 July 1975, no. A329/75, ALA, E, LANC. The plants were small and _ half hidden in a low bryophyte mat. The gully was only a metre ‘or so wide and deep. It drained a small flush, colorful with Primula tschuktschorum, and, in addition to the Minuartia, harbored Ranunculus eschscholtzii, Epilobium angallidifolium, and Anten- naria monocephala subsp. monocephala. THE CANADIAN FIELD-NATURALIST Vol. 91 This new locality is the only one between those in the Alaska Range and in the Brooks Range. Arenaria chamissonis Maguire Sourdough Creek: exposed ridge north-east of the head of the creek and south of Hope Mountain, 1150 m (3500’). 16 July 1975, no. A357/75, ALA, LANC. Only a few cushions of this inconspicuous, Cherleria-like plant were found. Arenaria chamissonis is almost restricted to Alaska, being previously known only from three areas — the western Brooks Range to the Bering Straits, the Seward and Chukotsky Peninsulas, and the Alaska Range. Ranunculus eschscholtzii Schlecht. Sourdough Creek: in snow-patch gully at head of creek, 980 m (3000’). 15 July 1975, no. A331/75, ALA, BM, E, LANC. This locality represents an extension northwards across the Tanana River from its previous limit in the Mount McKinley area. Potentilla rubricaulis Lehm. Galbraith Lake: north side of valley north-east of Pump Station 4. 19 July 1975, no. A429/75, ALA, LANC. The site was a dry, sheltered, south-facing slope below a small limestone cliff. The rich flora included Agropyron violaceum subsp. andinum, Bromus pumpellianus, Polemonium boreale, Myo- sotis alpestris, and Crepis nana. In the field, the few plants were clearly distinct from P. pulchella, seen shortly before at Prudhoe Bay, the leaflets being wider and softer. The material was submitted to K. Jakobsen, of Copenhagen University, who confirmed the identification, as also did Professor Hultén. Material from Galbraith Lake and east Greenland of this often misunderstood species and of P. pulchella from Prudhoe Bay and east Greenland are shown in Plate 1. This is the first record of P. rubricaulis from Alaska. Like Pleuropogon sabinei, it was confidently included by Hultén (1968) in his Alaskan Flora although at that time it was not known west of the Mackenzie River, Northwest Territories. Oxytropis deflexa (Pall.) DC. var. sericea Torr. & Gray Fairbanks: roadside gravel opposite the airport entrance. 2 July 1975, no. A25/75, ALA, LANC, S. The material is typical of this variety. The illus- trations of var. sericea and var. foliolosa (Hook.) Barneby in Hultén’s (1968) Flora are reversed; the latter has not been recorded from central Alaska. This find represents an extension of range north- westwards down the Tanana valley from Tanacross. Douglasia gormanii Constance Livengood: east-west ridge north of the Elliott Highway, 55 km south-west of Livengood, 730 m 1977 NOTES 321 FiGurE 1. Potentilla pulchella R.Br.: A — @rsted Dal, central east Greenland, G. Halliday, no. G68/74, 1974, C. LANC; C — Prudhoe Bay, Alaska, G. Halliday, no. A266/75, 1975, LANC. Potentilla rubricaulis Lehm.; B — Mestersvig, central east Greenland, G. Halliday, no. G20/74, 1974, C, LANC; D — Galbraith Lake, Alaska, G. Halliday, no. A429/75, 1975, ALA, LANC. (2400’). 4 July 1975, no. A94/75, ALA, LANC. Onlya few plants were found, well past flowering, growing in open, stony ground on the south side of the ridge and associated with Draba fladnizensis and Ligusticum mutellinoides. The map in Hultén’s (1968) Flora shows two areas for this rare Alaskan-Yukon endemic. The new locality effectively bridges the gap between them. Euphrasia disjuncta Fern. & Wieg. Sourdough Creek: track near junction of Sourdough and Polar Creeks, above the abandoned Placer Mine. 17 July 1975, no. A379/75, LANC. A few plants were found growing in the grassy center of the old overgrown track. This locality, Gyjaerevoll’s (1967) record from College, and a 1976 find by the English botanist E. C. Wallace (herb.E.C.W.) at Goldstream Crossing, 15 km west of Fairbanks, help to bridge the gap shown in Hultén (1968) between the Alaska Range and two isolated occurrences to the north, one on the Seward Peninsula, the other on the south side of the Brooks Range. Pedicularis hirsuta L. Prudhoe Bay: between the dock and the headland on the west side of the Sagavanirktok River. 11 July 1975, no. A216/75, ALA, LANC, S. Several plants were seen scattered along a terrace above the shore and occurring chiefly with Eriophorum triste and Carex stans in shallow depressions in an otherwise dry Dryas polygon vegetation. 322 This is the first record for this species in Alaska. Hultén (1958) regarded it asanamphi-Atlantic species with a distribution extending from near Coppermine, Northwest Territories (112°W) eastwards to the New Siberian Islands (140° E). The Alaskan locality (148° W) suggests that, like Pleuropogon sabinéi, it is best regarded as circumpolar. Antennaria monocephala DC. subsp. monocephala Sourdough Creek: in snow-patch gully at head of creek, 980 m (3000’). 15 July 1975, no. A322/75, LANC. The specimens were densely tufted and quite distinct from the stoloniferous subsp. philonipha (Pors.) Hult. which was found a few kilometres north- west of this site. Although this is the first montane record north of the Tanana River, it extends up to the coast to Cape Lisburne. In their provisional checklist of the vascular, bryophyte and lichen flora of Prudhoe Bay, Murray and Murray (1975) listed 150 species of vascular plants. This list can now be supplemented by the following 12 species (Herbarium specimens of all of these are in LANC.): Phippsia algida(Soland.) R. Br., Pleuropogon sabinei R.Br., Puccinellia angustata (R.Br.) Rand. & Redf., Colpodium vahlianum (Liebm.) Nevski, Ranunculus pallasii Schlecht., R. hyperboreus Rottb., Saxifraga rivularis L., S. foliolosa R.Br., Epilobium davuricum Fisch. var. arcticum (Sam.) Polunin, Polemonium acutifolium Willd., Pedicularis hirsuta L., Chrysanthemum bipinnatum L. subsp. bipinnatum. THE CANADIAN FIELD-NATURALIST Vol. 91 Acknowledgments I am deeply grateful to D. F. Murray, University Museum, University of Alaska; the Institute of Arctic Biology, University of Alaska; B. P. Alaska Inc. and Alyeska Inc. for invaluable support in Alaska; and to D. F. Murray and E. Hultén for assistance in the preparation of this note. Literature Cited Gjaerevoll, O. 1967. Botanical investigations in central Alaska, especially in White Mts, 3. Det Kongelige Norske Videnskabers Selskabs Skrifter 1967, 10. Hultén, E. 1958. The amphi-Atlantic plants and their phytogeographical connections. Kungliga Svenska Vetenskapsakademiens Handlingar, Series 4, 7(1). Hultén, E. 1968. Flora of Alaska and neighboring ter- ritories. Stanford University Press, Stanford. 1008 pp. - Murray, B.M. and D.F. Murray. 1975. Appendix A: Provisional checklist to the vascular, bryophyte, and lichen flora of Prudhoe Bay, Alaska. Jn Ecological investigations of the tundra biome in the Prudhoe Bay Region, Alaska. Edited by J. Brown. Biological Paper of the University of Alaska, Special Report Number 2. pp. 203-212. Scotter, G. W. and W. J. Cody. 1974. Vascular plants of ‘Nahanni National Park and vicinity, Northwest Ter- ritories. Naturaliste Canadien 101: 861-891. Tzvelev, N.N. 1964. Puccinellia Parl. In Arkitcheskaya Flora SSSR 2. Edited by A. 1. Tolmachev. Academy of Sciences of the SSSR, Komarov Botanical Institute, Moscow and Leningrad. pp. 178-208. Received 18 August 1976 Accepted 13 July 1977 Range Extensions of the Water Shrew and Mink Frog in the James Bay Region of Quebec D. J. OXLEY, R. A. CouTTs, and N. G. H. BOYLE Department of Biology, Carleton University, Ottawa, Ontario KIS 5B6 Oxley, D.J., R.A. Coutts, and N.G.H. Boyle. 1977. Range extensions of the Water Shrew and Mink Frog in the James Bay region of Quebec. Canadian Field-Naturalist 91(3): 322-323. Between 17 June and 28 August 1973 we observed Mink Frogs (Rana septentrionalis) and Water Shrews (Sorex palustris) in the vicinity of Kanaaupscow Post (54°05’ N, 76°29’ W). Mink Frogs have been collected at Lac Aigneau (Logier and Toner 1961), Lac Nathalie (MacCulloch and Bider 1975) and further east in Ungava (Conant 1975). During our survey, three adult Mink Frogs were sighted, two of which were captured. The coloration of these specimens was variable; one frog had an irregularly mottled pattern and the other had distinct spots. Although Wood Frogs (Rana sylvatica) and American Toads (Bufo americanus) were common, Mink Frogs were rare. The three sightings occurred only on small creeks; these had a mud and detritus bottom, stabilized by beaver dams, and the predominant vegetation was lilies, grasses, sedges, and dwarf willows. Our specimens, representing a range extension of 80 km in northwestern Quebec, have been catalogued at the National Museum of 1977 Natural Sciences as NMNS 17492 and 17493; they were collected 31 July and 15 August 1973. Both were mature females with developing eggs and measured 70.5 and 65.0 mm after preservation. Their large size agrees with similar measurements found for other northern Mink Frog populations by Schueler (1975). The two adult female Water Shrews were found on shore within 2 m of the Kanaaupscow River. One specimen was taken in a Museum Special trap in sedge-willow scrub. The other shrew was found dead on a sandy beach and large canid tracks around the body along with injuries on the specimen indicated that it had probably been killed and discarded by a wolf (Canis lupus). The previous known occurrences of the Water Shrew were recorded on the opposite side of James Bay at Cape Henrietta Maria and further north in Ungava at Fort Chimo (Peterson 1966). Our records represent an extension of over 300 km for the shrew in northwestern Quebec. The specimens have been catalogued at the Carleton University Museum of Zoology as CUMZ 4876 and 4877; total length measured 168 mm and 166 mn, tail vertebrae 80 mm and 80 mm, and weight 10.6 and 9.9 g. We thank Francis R. Cook of the National Museum of Natural Sciences for confirming identifi- NOTES 328 cation of the Mink Frogs, and C. G. van Zyll de Jong of the National Museum for providing distributional data on the Water Shrew. We especially thank M. B. Fenton of Carleton University for his help in all aspects of the work. This study was supported by a grant from the Indian and Inuit Association of Northern Quebec. Literature Cited Conant, R. 1975. A field guide to reptiles and amphibians of eastern and central North America. Houghton Mifflin Company, Boston. xvili-429 pp. Logier, E. B.S. and G. C. Toner. 1961. Check list of the amphibians and reptiles of Canada and Alaska. Royal Ontario Museum, Life Sciences, Contribution 53: 1-92. MacCulloch, R. D. and J. R. Bider. 1975. New records of amphibians and garter snakes in the James Bay area of Quebec. Canadian Field-Naturalist 89: 80-82. Peterson, R.L. 1966. The mammals of eastern Canada. University of Toronto Press, Toronto. 465 pp. Schueler, F. W. 1975. Geographic variation in the size of Rana septentrionalis in Quebec, Ontario and Manitoba. Journal of Herpetology 9: 177-185. Received 30 April 1976 Accepted 12 March 1977 Range of the Bushy-tailed Wood Rat (Neotoma cinerea) in Alberta DAVID W. KRAUSE! and BRUCE G. NAYLOR?2 ‘Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9 2Department of Geology, University of Alberta, Edmonton, Alberta T6G 2E3 Krause, David W. and Bruce G. Naylor. 1977. Canadian Field-Naturalist 91(3): 323-324. Soper (1961, p. 34) has stated that the Bushy-tailed Wood Rat (Neotoma cinerea) “is essentially a dweller in the Rocky Mountains, but stragglers infiltrate suitable Great Plains environment to a very limited extent.” Elsewhere, Soper (1964, p. 199) noted an “unusual” record of this species from Ponoka, in central Alberta. Evidence presented here documents additional occurrences of the Bushy-tailed Wood Rat on the plains of Alberta. A complete articulated skeleton (University of Alberta Museum of Zoology catalogue number 7769) of N. cinerea drummondii was discovered in September of 1973 along the Smoky River some 35 km east of Grande Prairie (NW 1/4, Sect. 16, Twp. 72, Rg. 2, W. 6). The unbleached state of the bone indicates a recent death and the articulated Range of the Bushy-tailed Wood Rat (Neotoma cinerea) in Alberta. condition makes transport by predator unlikely. In July 1974 a nearly complete skeleton (UAMZ 7770) of N. c. cinerea, with skull and jaws intact, was recovered approximately 27 km south of Empress, Alberta at the Trans-Canada pipeline crossing of the South Saskatchewan River (NW 1/4, Sect. 18, Twp. 20, Rg. 1, W. 4). At the same time, the anterior part of a second skull (UAMZ 7771), withan associated left dentary, was found some 100 m south- east of the more complete find. These two specimens constitute a range extension of approximately 150 km north of previously known occurrences along the Milk River (Soper 1946; Rand 1948) and its tribu- taries (Nero 1956). Another, although questionable, range extension 1s a record from the town of Strathmore, some 61 km 324 Y N. c. drummondii a N. c. cinerea 0 100mi 0 100km FIGURE |. The range of the wood rat in Alberta. Hatching and cross-hatching indicate the well-established ranges of Neotoma cinerea drummondii and N. c. cinerea, respectively (after Soper 1964). Extralimital occurrences are depicted by the following symbols: , Brownyale (Kelsall 1971); A, Ponoka (Soper 1964); O:, Smoky River; O2, Strathmore; O3;, Empress. Abbreviations designate major urban centers. east of Calgary. This specimen was apparently collected by W. Sturm, a resident of Strathmore, in April 1970 and was subsequently donated to the Provincial Museum and Archives of Alberta, Edmonton (catalogue number Z72.102.26). More precise locality data are not available. Previous extralimital extensions for the Bushy- tailed Wood Rat east of the Canadian Rocky THE CANADIAN FIELD-NATURALIST fe) ° S Nn = Vol. 91 Mountains have been reported by Nero (1956) at Govenlock, Saskatchewan (in the extreme south- western corner of the province), Soper (1964) at Ponoka, and Kelsall (1971) at Brownvale (near the town of Peace River, Alberta). The species appears to be established on the prairies in areas in, or adjacent to, major river valleys proximate to the Rockies. The records from Govenlock, Ponoka, Brownvale, Smoky River, Empress, and perhaps Strathmore document occurrences near the Milk, Battle, Peace, Smoky, South Saskatchewan, and Bow Rivers, respectively. The rocky exposures found along these rivers provide suitable habitat for the wood rat on the prairies. In those instances where the Bushy-tailed Wood Rat has been captured some distance from a river valley, it has been found inhabiting an occupied or abandoned human dwelling, a habit that is common for the wood rat (Rand 1948; Banfield 1974). It would be of considerable interest to establish the extent of the range of the wood rat on the prairie areas of Western Canada by a collection program along the major river valleys. Acknowledgments We thank N. Panter, former Curator, and W. E. Roberts, present Curator of the Museum of Zoology, The University of Alberta, and H. Smith, Curator of Mammals, Provincial Museum and Archives of Alberta, Edmonton for permitting access to speci- mens in their charge. Literature Cited Banfield, A. W. F. 1974. The mammals of Canada. Univer- sity of Toronto Press, Toronto. xxv + 438 pp. Kelsall, J. P. 1971. A range extension for the bushy-tailed wood rat. Canadian Field-Naturalist 85(4): 326. Nero, R. W. 1956. A record of the packrat in Saskatch- ewan. Blue Jay 14(2): 43-44. Rand, A. L. 1948. Mammals of the eastern Rockies and western plains of Canada. National Museum of Canada Bulletin 108. 237 pp. Soper, J. D. 1946. Mammals of the northern Great Plains along the international boundary in Canada. Journal of Mammalogy 27(2): 127-153. Soper, J.D. 1961. Field data on the mammals of southern Saskatchewan. Canadian Field-Naturalist 75(1): 23-42. Soper, J.D. 1964. The mammals of Alberta. Queen’s Printer, Edmonton. 402 pp. Received 20 January 1977 Accepted 23 March 1977 News and Comment Notice of Change to the By-laws of The Ottawa Field-Naturalists’ Club Changes to By-laws 12, 15, and 16 of The Ottawa Field-Naturalists’ Club were passed unanimously by the Council at the meeting of 2 May 1977. These By-laws now read as follows: 12. Disbursements of Club Monies Disbursements of Club monies shall be made by the Treasurer on receipt of properly rendered accounts verified by the Chairman of the Com- mittee concerned or by a Business Manager or as specified by the Council. Disbursements of Club monies shall be made only by cheque bearing the signature of any one of the three following members of the Council: President, Treasurer, Business Manager of The Canadian Field-Naturalist. 15. Annual Dues The schedule of annual dues shall be as follows: Memberships — Individual $10.00 Family $12.00 16. Subscription Fees The schedule of subscription fees shall be as follows: The Canadian Field- Naturalist — Individual $10.00 Libraries and Institutions $20.00 Trail and Landscape — Libraries and Institutions $10.00 Diana R. Laubitz, Recording Secretary Wanted: Data on the Seasonal Distribution of North American Gulls We are developing a procedure whereby the U.S. Air Force can predict the potential seasonal hazard to aircraft represented by gulls in parts of North America. This knowledge will be used to schedule missions around high-risk areas, thereby reducing the likelihood of bird/aircraft collisions. Supplemental data on local gull populations are needed from all parts of the continent. The assistance of field workers is solicited to aid us in this task. Please submit reports of your gull observations to Dr. William E. Southern, Department of Biological Sciences, Northern Illinois Symposium on Wapiti A symposium on the ecology and management of wapiti with particular reference to the Jackson Hole- Yellowstone ecosystem is planned for 3 to 5 April 1978. It will be hosted by the Department of Zoology and Physiology at the University of Wyoming. Papers are welcome onall aspects of the biology, ecology, and University, DeKalb, Illinois 60115. Data will be gathered for a 2-year period beginning | September 1977. For each observation, please provide the following information: list of species present, approximate number of each species, precise locality description, dates observed, any information about causes for concentrations (e.g., sanitary landfill operation), and any details about the frequency of such concen- trations in the respective areas. Information is sought from inland as well as coastal localities. management of wapiti (elk) and are not restricted to research projects conducted specifically in the Jackson-Yellowstone region. Titles and abstracts should be sent by 31 December 1977 to Dr. M. Boyce, Box 3166 Univ. Sta., Laramie, Wyoming 82071. 325 Book Review ZOOLOGY Biology of Insects By David J. Horn. 1976. Saunders, vil + 439 pp. $14.50. Philadelphia. This book was written to fill “a need for a basic text that regards insects as adaptive solutions to problems of survival in a heterogeneous and largely hostile environment.” It seeks “to show how insects function and how selective pressures have shaped [their diversity].” The idea is very good; unfortunately its execution leaves much to be desired. Though Dr. Horn’s warm and human introduction disposes one favorably to his efforts, and though he has dealt witha wide range of subjects and included a wealth of interesting information, his book suffers from several major faults as well as from a large number of imperfections of detail. First, it is hard to imagine what class of readers the book is meant for. Though the author is a university teacher, the book is written at a conceptual level better suited to a high school. The language, too, is high- school English, subspecies nearcticus. On the other hand, what high-school course would accommodatea 400-page text on entomology? What student ready for an entomology course needs a grossly oversimplified account of the scientific method as an introduction? Who needs the elementary ideas of the biological species concept or of the origin of adaptation by natural selection explained, yet is ready for r- and K- strategies, for life tables, or for integrated control? Why give 120 pages to a non-diagnostic and hardly descriptive survey of insect orders and families? Second, the text has not been well thought through. It is an inventory rather than a synthesis. Topics tend to be related by cross-reference rather than by integrated discussion. Difficult subjects are dismissed rather than dealt with. Numerical taxonomy is characterized in six lines, in which it is said to “have become a valuable addition to the systematic tool- box.” Nutrition gets one page, indicating that insects need proteins, amino acids, carbohydrates, sterols, vitamins, mineral ions, and water. Terminology often is emphasized at the expense of concepts, and the emphasis given to terms seems rather erratic. For example, in the two pages on “Integument,” the following terms are given in bold face: “chitin,” “sclerotin,” “apodemes,” and “tentorium.” A number of other equally important terms are given in ordinary type or omitted. The same lack of finish extends down to paragraphs and sentences: parentheses are often used to bolster a passage that should have been rewritten, e.g., “Most spiders feed on insects (and other spiders) though some of the largest tropical species can (and do) eat small fish and birds.” A third major defect is the poor quality of the illustrations. Most of the drawings are crude, many are inaccurate or unrecognizable, such as the clothes moth in Figure 3-55 and the gelechiid in Figure 3-57. The caterpillars shown in the illustration of holo- metabolous development in Figure 2-13 have prolegs on every abdominal segment except usually the anal, and appear to add abdominal segments as they grow. The three supposed Colias species in Figure 4-26 bear no resemblance to any Colias | have ever seen, but might be inaccurate renditions of cabbage butterflies. A large proportion of the photographic illustrations are also unsatisfactory, being out of focus, low in contrast, based on damaged material, or otherwise substandard. On a more detailed level the errors and incon- sistencies are too numerous for individual mention. The International Commission on Zoological Nomenclature makes, interprets, and at times sus- pends rules, but does not police them as stated on p. 13. Ordinal, family, and other higher-group names are sometimes used as singular nouns, sometimes as plural. They should be plural. Antheraea is con- sistently misspelled Anthraea. The armyworm is Pseudaletia unipuncta, not unipunctata, as given on p. 119. On the same page and elsewhere the plural abbreviation “spp.” is used in place of the singular “sp.” Figure 3-60 A, labelled “Indian meal moth, family Pyralidae,” is a microlepidopteran, not an Indian meal moth or a pyralid. I think it only fair to say that in reading Biology of Insects 1 learned many things about insects and their biology that I was unaware of before. The biblio- graphy is extensive and includes a good selection of useful titles. Here, too, however, caution is needed. Holland’s Moth Book was first published in 1903, not 1913. The reference to Dominick et al., The Moths of America North of Mexico, contains five separate errors of citation. I regret that for most readers Biology of Insects cannot be recommended. EUGENE MUNROE Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6 326 1977 BOOK REVIEWS 327 North American Game Birds of Upland and Shoreline By Paul A. Johnsgard. 1975. Press, Lincoln. 183 pp. $6.95. University of Nebraska North American Game Birds of Upland and Shoreline “is intended as a guide to the more common species of American game birds.” Johnsgard “decided to include all of the native and successfully intro- duced species of gallinaceous game birds of North America occurring north of Mexico as well as the migratory game birds other than waterfowl that are legally harvested in significant numbers in the United States and Canada.” A total of 29 species, organized by order, are listed in the book. The treatment given each species is standardized. For each species there are sections on distributional range, identification, field marks, age and sex criteria, habitat and foods, social behavior, and reproductive biology. Line drawings, black-and-white and color photographs are also included within the book, although not all species are equally treated. Within the section on reproductive biology the material contained includes information on egg- laying, clutch size, incubation period, parental care and protection, and time to flight. The section on social behavior outlines the species’ territorality and flocking behavior. The age and sex criteria section contains information on how to distinguish between males and females of the species as well as how to identify immature birds from adults. Under field Waterfowl Studies By Bruce Burk. 1976. Winchester Press, New York. (Canadian distributer Nelson, Foster and Scott, Willow- dale). 254 pp., illus. $17.95. “The availability of good wildlife photographs has played by far the most important role in the development of wildlife art.” With this thought in mind, Waterfowl Studies by Bruce Burk was intended to assist the wildlife artist. The book provides a reference collection of waterfowl photographs “for the decoy maker and collector, hunter, bird watcher, waterfowl artist and naturalist.” “The text of this book has been kept to a minimum to yprovide maximum space for photographs.” Burk’s collection of North American waterfowl photographs include 21 ducks, 8 geese, and 2 swans. Each species is photographed from a variety of angles so as to record the characteristic shapes and features of the species profiled. “For the first time in one volume, body form, position and plumage detail are accurately recorded.” Waterfowl Studies does not include the various marks, Johnsgard points out some of the seasonal variations that occur within the species as well as the features that distinguish one species from another similar-looking species. Generally speaking the book is very well organized and contains a great deal of material in capsulized form. Johnsgard includes in his book a very interesting section entitled ‘The Hunting and Recreational Value of Upland Game Birds.’ Within this section he points out that approximately a billion dollars annually is spent in pursuit of small game. Furthermore “the average hunter of upland game birds might be expected to kill in the course of a season about 7 birds.” No matter how important hunting appears to be, “it is clear that the proportion of Americans who elect to enjoy wildlife through non-consumptive methods may be at least as great as or possibly greater than those who prefer to carry a gun when afield.” Johnsgard’s book is highly recommended to all persons wishing to know a little more about game birds. As the author himself states “the book is intended to be fuller than the typical bare-bones field guides but less detailed than a full scale monographic treatment.” PETER CROSKERY Ontario Ministry of Natural Resources, Ignace, Ontario marine ducks of North America, it concentrates on the inland species. For some reason Burk has included only one of the three common merganser species, the Hooded Merganser. Omitted are the American Merganser and the Red-breasted Merganser. About 90% of the included photographs are in black and white. Some of these black-and-white photographs are grainy, probably the result of the film type and speed preferred by the author. Nearly all included photographs are close-up pictures of the birds profiled. As. Burk states “although this collection of water- fowl photos has been assembled primarily to aid the artist in making realistic bird carvings and paintings, these pictures will also be of interest to other bird lovers and naturalists.” One could not really consider this book as informative; however I found it very enjoyable. PETER CROSKERY Ontario Ministry of Natural Resources, Ignace, Ontario 328 An American Crusade for Wildlife By J. B. Trefethen. 1975. Wincester Press and Boone and Crockett Club Book, New York. 383 pp. $14.50. An American Crusade For Wildlife is a historical review of wildlife conservation in America. The period covered by the review 1s from the arrival of the first Europeans in North America until the present day. Within the text, Trefethen reports the factual history of wildlife conservation, but more importantly he reveals the changes in wildlife conservation philosophy that have evolved in North America. In the days of the early settlers the wildlife resources of North America were never considered as a limited resource. They were simply there and were to be utilized at the imagination of the user. Under such an attitude evolved the buffalo hunter, the mountain man, and the market hunter. Primarily as a result of the activities of these commercial resource consumers, the limits to wildlife resources became apparent. Buffalo numbers dwindled, furbearers such as beaver declined, and deer vanished from parts of North America. The North American society of the day saw the end nearing for species previously abundant. Another interesting aspect of wildlife conservation, as pointed out (indirectly) by Trefethen, is that where change was necessary, the proposal for change generally originated from outside the political realm. Most of the early legislation regarding wildlife conservation was proposed, lobbied for, and sup- ported by, public conservation action groups. An American Crusade for Wildlife reviews the origin and original purposes for some of the groups we know today. America’s Master of Bee Culture. The Life of L. By Florence Nail. 1976. Cornell University Press, Ithaca N.Y. 215 pp. $9.95. There are few industries which can pinpoint the day when the major discovery was made affecting their future development. Florence Nail, in her book, “The Life of L. L. Langstroth,” has done just that. By careful research of all of the writings of Langstroth and his personal notes, Florence Nail has been able to pinpoint this as October 30th, 1851, and she has even been able to reproduce his drawings of that date. Langstroth’s discovery was based on the fact that the bees use space of approximately 3/8 inch through which to pass and they will not fill this space either with bee glue (propolis) or wax. Langstroth stated on the day of his discovery, “The use of these frames will, I am persuaded, give a new impetus to the easy and THE CANADIAN FIELD-NATURALIST Vol. 91 Wildlife resources have seldom been a subject of political party platforms. An American Crusade for Wildlife suggests that through history individual politicians, regardless of party, have proposed or supported wildlife conservation efforts. It would appear that even among politicans, wildlife conser- vation 1s a very personal subject. The pressures upon wildlife resources have resulted primarily from human population increases. “The heaviest impact of the white settlement came not so much from direct killing that attended it as from the wrenching changes in the prevailing habitat brought about by the intrusion of European farming.” No book on the wildlife conservation movement would be complete without comments on today’s issues and problems. If An American Crusade for Wildlife has a weak point, then this would be it. Trefethen’s list of topics of concern of today’s wildlife conservationists is rather short and incom- plete. It includes endangered species, seal hunting, and oil spills. It does not include environmental education, resource management of private land, or user group conflicts. An American Crusade for Wildlife is very much United States oriented; even so, it makes fascinating reading. One should remember that many events in United States conservation history have had direct parallels in Canada. PETER CROSKERY Ontario Ministry of Natural Resources, Ignace, Ontario L. Langstroth profitable management of bees.” Little did he realize how true these words were. Florence Nail, in a very interesting manner, has portrayed not only all of the factors leading up to this discovery but the trials and tribulations through which Langstroth went during his lifetime’s work; he was a minister of the Christian gospel, the inventor of the first practical moveable frame beehive, and author of a book, “The Hive and the Honeybee,” which will forever remain a classic in bee culture. “The Hive and the Honeybee,’ now published in an absolutely different form and recently extensively revised in 1975, is a fitting tribute to the original author and is still recognized as the beekeepers’ leading manual. Florence Nail’s account of Langstroth’s life and work is more than a biography of L. L. Langstroth; it NOFA portrays the struggle to improve the beehive and the fact that even though Langstroth realized the im- portance of his discovery, many of those who made use of it, did not until much later in time. Florence Nail was influenced, to a great extent, in her preparation of the manuscript, by Dr. E. F. Phillips, who must have been a great inspiration to her as, at that time, he was one of the leaders of the industry. Those who knew Dr. Phillips would be interested in the introduction which he wrote and which portrays rather clearly the controversies of even that period, of the late ’30s and early ’40s, which were going on BOOK REVIEWS 329 between Phillips and other leaders of the industry. He mentions fads such as top entrances in winter preparation, a practice which has now become quite common in areas where wintering is difficult. This fascinating book, by Florence Nail, is a must for all interested in apiculture, whether they be scientists, hobbyists, or commercial producers. G. F. TOWNSEND Department of Environmental Biology, University of Guelph, Ontario NIG 2WI Biology of the Kaminuriak Population of Barren-ground Caribou. Part 4: Growth, Reproduction and Energy Reserves By T. C. Dauphiné, Jr. 1976. Canadian Wildlife Service, Ottawa. Report Series 32. 71 pp. $3.25. This is the fourth and final report dealing with the intensive study of the Kaminuriak caribou population conducted by biologists of the Canadian Wildlife Service in cooperation with the governments of the Northwest Territories, Manitoba, and Saskatchewan, between 1966 and 1968. Earlier parts by G. R. Parker (1974), F. L. Miller (1974), and D.R. Miller (1976) were published in the same report series. The present study relies on conclusions reached in the previous parts with reference to population, age, and diet, to analyze the anatomical data obtained from post mortem examinations. In some respects it is the most important of the parts, in that conclusions were drawn on various environmental factors that are controlling the population size. Basic management procedures are recommended to maintain the population as well. A number of vital statistics dealing with growth, reproduction, and survival of this population have been determined. In comparison with other mainland populations, the Kaminuriak caribou showed slow growth, modest size, and delayed puberty, which is thought to be a reflection on their environment. Perhaps the most interesting findings were related to the annual nutritional and reproductive cycles. Increases in body size and weight were restricted to a summer growth period from June to October. Mature caribou were heaviest in autumn immediately prior to the rut. Adult males lost about 30 percent of their maximum weight during the rut, but managed to maintain that weight until the next growth cycle began in spring. Adult females lost 11 percent of their mean autumn weight by late winter and did not begin to regain weight until mid-July. This annual cycle of weight change was confirmed in the size of fat deposits. Both sexes had maximum fat reserves in September. The mature males mobil- ized these reserves during the rut when the back and visceral deposits were rapidly depleted. Pregnant females maintained high fat reserves in the autumn, but experienced a steady decline of reserves during the winter and spring, to reach their nadir in June and early July at the time of parturition and lactation. A higher percentage of females than males mobilized marrow fat reserves. Over half the 1'4-year-old females ovulated but only 2 percent conceived; 48 percent conceived at the age of 2'4 years and 90 percent conceived at 34 years and older. Ovarian scars indicated that an average cow bore a calf four out of every five years. Thin cows generally speaking did not conceive. The observations suggest a cycle of gradual exhaustion of body condition during successive pregnancies, followed by a barren year accompanied by physical recuperation and then renewed breeding activity. Calf production was generally high, but depth of winter snow seemed to be an important factor in calf survival to the yearling class. The findings of this study indicate the key impor- tance of summer range to the survival of Kaminuriak caribou. This was especially true of the females which reached their nadir of stored fat at the time they were nursing their calves. The amount of fat they canamass by autumn appears to influence their chances to conceive and the survival of calves. Indicated management strategies included the harvesting of a greater percentage of males, and possibly wolf control. It was also recommended that human disturbance of caribou on the summer range should be carefully regulated. A. W. F. BANFIELD Institute of Urban and Environmental Studies, Brock University, St. Catharines, Ontario L2S 3A] 330 THE CANADIAN FIELD-NATURALIST Vol. 91 Répartition géographique des poissons, leur abondance relative et bathymétrie de la région du Lac Saint-Pierre By Gérard Massé et Jean-René Mongeau. 1974. Service de Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. 59 pp., illus. Cartes de répartition géographique des espéces de poissons au sud du Québec, d’aprés les inventaires ichthyologiques effectués de 1963 a 1972 By Jean-René Mongeau, Albert Courtemanche, Gérard Massé, et Bernard Vicent. 1974. Faune du Québec, Rapport Spécial (4): 1-92. Les poissons de la région de Montréal, la péche sportive et commerciale, les ensemencements, les frayéres, la contamination par le mercure et les PCB By Jean-René Mongeau et Gérard Massé. 1976. Service de Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. 286 pp., illus. Gaps in occurrence on species distribution maps frequently result from lack of collecting rather than absence of the species. Museums and environmental agencies frequently lack the funding or planning that would result in thorough zoological and botanical surveys that would produce excellent spot distribu- tion maps. It is desirable that such maps be based on voucher specimens whose identity can be verified when called in doubt or when taxonomic changes are made. High-quality distribution maps serve a number of useful purposes including the following: (1) recreational use — where to go to bird-watch or fish, (2) resource use — where to go to catch fish, (3) resource management, (4) determination of effects of environmental distur- bance or pollution, (5) zoogeographic studies, (6) ecological studies. Méthodes de péche expérimentale, en eau douce, a By Jean-René Mongeau. 1976. Rapport Technique, Service de lAménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche. 83 pp. Biologists and technicians trained in universities receive too little training in the use and care of fishing gear, even though it is very important in sampling populations and in the collection of specimens. Under one cover Mongeau has brought together information on techniques of fishing with gill nets (including setting them under ice), seines, traps, trap nets, dip Since 1963 the Ministére du Tourisme, de la Chasse et de la Péche has been surveying the fish fauna of southern Quebec. Samples have been made every 1/5 mile along water courses. Such methodical sampling with a variety of gear permits the drafting of distribution maps which indeed show where a species is present and where it is absent. The appearance of the results has begun in the above publications. One species is presented per map with the stations positive for the species indicated bya dark spot. Some maps even indicate whether the species was rare (<50), abundant (50-200), and very abundant (200). The ministry and its team of workers is to be commended for the production of these fine maps which will be useful to many groups. It is hoped that other organizations and museums will be given the resources to produce similar atlases for our fauna and flora. DON E. MCALLISTER National Museum of Natural Sciences, Ottawa, Ontario KIA 0M8 Pusage du biologiste et du technicien de la faune nets, set lines, electric shockers, and rotenone. De- tailed specifications are given for gill nets and seines and instructions on care are offered. The text and photos together offer an excellent introduction to the techniques of fishing. DON E. McALLISTER National Museum of Natural Sciences, Ottawa, Ontario KIA OM8& 1977 The Anatomy of Fishes. Parts I and II. By Wilhelm Harder. 1975. E. Schweizerbart’sche Ver- lagsbuchhandlung, Stuttgart. Part I, 612 pp. Part II, 132 pp., 337 figures, 13 plates. $96.40 (U.S.). Since Harmer’s (1904) The Cambridge natural history and Goodrich’s (1909) Cyclostomes and fishes, there has not been an in-depth treatment of the anatomy of fishes in the English language. There is a need, because fishes are an ancient group, rich in taxa which are more highly diversified in anatomical structure than other more recent vertebrate classes. Harder’s Anatomy of fishes, a translation and revision of an earlier German text, admirably fills the lacuna. My first impression, one never dislodged, was that here was a source book, solidly packed with infor- mation. Closer inspection suggested that even the machete of today’s journal editor would go rusty whilst seeking verbiage to slash. A fair amount of text is devoted to basic zoology (definition of positional terms such as caudad, discussion of protoplasm and cells) or science (resolv- ing power of light microscopes). This means that the text could be used without difficulty by beginning university students. But the text then proceeds to deeper levels, and more detail at the cellular and tissue level is given than in the usual ichthyology text. Anatomical terms are in italics, making it easy to skim through paragraphs for terminology. The scientific names of fishes are in ordinary type face. A stream of functional thought runs through the chapters; it is not just a descriptive anatomy. The references are grouped, to the reviewer’s mind inconveniently, by chapter at the end of the book. This means that first the chapter group must be sought before finding the references. The reviewer finds the level of scientific accuracy high and the breadth coverage very good, although the author modestly admits the impossibility of complete coverage. A few lapsi may be indicated. It is stated that the radii branchiostegi develop phylo- genetically out of the lateral gularia (it is, rather, the reverse), also, that in general there is only one dorsal Wolf... Kill! The Wilderness Called Shunka By Marika Lumi. 1976. Van Toronto. 195 pp., illus. $9.95. Nostrand Reinhold, The title Wolf... Kill! was presumably chosen to ensure this book sells well. It is somewhat ambiguous, however, because the hero, tundra wolf Shunka, is anything but a killer. No doubt the title is meant to refer to many people’s attitude toward the wolf: it should be shot on sight. The whole point of the book is BOOK REVIEWS 331 fin-bearing element per muscle-segment (usually, in fact, there is more than one, the prime exception being the blennioids). D. Rosen’s work on cephalic muscles in fishes was not cited. According to V. M. Maku- shok, contrary to Harder, trunk lateral lines are present in almost all fishes reported to be lacking them, including clupeids; it is only the canal and pores that are missing. Two works worthy of mention published too late for Harder to include are these: R. Winterbottom (1974), A descriptive synonymy of the striated muscles of the Teleostei (Proceedings of the Academy of Natural Sciences Philadelphia 125(12): 225-317); and J. S. Nelson (1976), Fishes of the world (John Wiley and Sons, New York, London, Sydney, Toronto. 416 pp.). The latter would have provided an excellent basis for classification. ; The translator has generally done a good job grammatically. He has done less well with the terminology (fin-sails instead of fin lobes, under- water-floor instead of bottom, bristle-teeth instead of cardiform teeth, keel bone in preference to para- sphenoid, but such lapsi are not common). For the most part these will be passed over without problem by the reader. The editing has not been of high quality. Pages 4 to 6, for example, contained seven misspel- lings, grammatical infelicities or taxonomic errors. Throughout the volume one finds similar errors (Vladikov for Vladykov, Colette for Collette, diver- ticle instead of diverticulum). The greatest careless- ness is found in regard to the bibliography. In looking up references in five pages of text, seven are found to be missing, two had disagreements in dates between the text citations and the bibliography. Such careless- ness is not acceptable. Despite its editorial shortcomings Harder’s Ana- tomy of fishes is an excellent reference book. Don E. MCALLISTER Curator of Fishes, National Museum of Natural Sciences, Ottawa, Ontario KIA 0M8 that wolves can be brought up to live amicably with people, in this case often sharing their house, their food, and their leisure time. Marika Lumi and her husband wanted to prejudice people in favor of wolves, which they have been able to do with their films, this book, and by means of Shunka himself. Neither is the subtitle about wilderness entirely suitable, for although Shunka is supposed to typify 332 for us the wilderness, he was in fact bought as a cub from the Alberta Game Farm near Edmonton. The author and her husband then raised him for a year first in Toronto, and then in the country near Pickering. When they could no longer manage him they sent him to a Natural Science School near Ottawa where he thrilled hundreds of school children who were allowed to visit with him in his pen. Eventually he was sent to British Columbia where the people were less under- standing about wolves and where he was finally shot after he had escaped. The author and her husband became devoted to Shunka, Marika Lumi describing him so affection- ately that we come to admire him too. Although in general Shunka was treated as far as possible as a dog, and was given a dog as a constant companion, he never acted entirely like one. He loved to romp and play with human beings and to join with howls in their A Field Guide to Birds’ Nests By H.H. Harrison. 1975. Houghton Mifflin Company, Boston. 257 pp., illus. $10.75. This book was written as an accompanyment to “Field Guide to the Birds” by R. T. Peterson (1934. Houghton Mifflin Company, Boston) to enable identification of the nest of a particular species without seeing the adult birds. It describes common nesting sites, nest construction, and characteristics of the eggs of all species of birds in the United States east of the Mississippi River. The author notes that usefulness of the book diminishes in the Canadian boreal forest. The format includes a color photo- graph of the nest and eggs of most species, followed by sections describing breeding range, habitat, nest, and eggs. A further section entitled “Notes” presents facts or personal experiences of the author for each species. Excellent black-and-white sketches of most birds by Ned Smith are also included. Detail provided by the author’s photography is exceptional. Combined with the written description, a vivid portrayal of nest and eggs is attained. For example, the user can readily identify the nests of most species of sparrows through egg characteristics. Other species are much more difficult. Some nests, particu- larly those of the cavity nesters, were disturbed for photographing. In several cases, for example, Long- billed Marsh Wren, the photograph would have been more meaningful if the nest had not been opened. In others man-made nesting sites were used, detracting from the value of the picture. Minor errors or incomplete explanations are found in certain of the text. The Spotted Sandpiper is described in a misleading fashion as inhabiting open THE CANADIAN FIELD-NATURALIST Vol. 91 sing songs, but his owners were never able to housebreak him, nor to teach him not to chew up books, shoes, or anything else he could fit into his mouth. Shunka considered his owners not as dogs would, but as members of his pack. He was dominant to the dog Happy and to the author, but submissive to the family cat and to the man of the house who controlled his life, feeding him, handling him on the leash, taking burrs from his coat, and driving him to the vet. This book about Shunka gives us a good idea of the nature of a wolf and will, I hope, create an improved climate of opinion for this fascinating species. ANNE INNIS DAGG Box 747, Waterloo, Ontario N2J 4C2 prairies, dry fields, and: pastures. Contrary to the statement in the preface, not all ducks commonly lay in one anothers’ nests. Species that are char- acteristically parasitic should have been noted. The discussion of evolution of nests is somewhat teleological although this problem is difficult to avoid, particularly in a book of this nature. The Redhead is the only species incorrectly omitted, although the author notes others which might have been included. Items that may have enhanced this book include a description of nesting period (done for some birds but not all), and a section similar to that found in “Field Guide to the Birds” listing like nests and eggs of other species and any differences. Ability to use this book would be improved considerably through develop- ment of a key based on habitat and location of the nest. As it is, the user requires a good knowledge of bird species to locate readily the correct photograph. Also, a statement of the dangers of disturbing birds during the laying period, numerous successive visits, and handling of fragile passerine eggs would have been of value. On the whole, however, the text itself is informative, useful, and apparently free of editorial errors. Accomplished bird-watchers and beginners will find this book useful in some situations where the nest and eggs are particularly diagnostic of the species. But, with few exceptions, the only way to be absolutely certain in identification of a nest remains in seeing the adult. IAN D. THOMPSON P.O. Box 895, Cochrane, Ontario POL 1C0 NOT BOOK REVIEWS 333) Pictorial Guide to the Mammals of North America By Leonard Lee Rue III. 1967. Apollo Editions, Crowell, New York. 299 pp., illus. $9.95. This attractive book is more than a pictorial guide; it has two or three pages of text devoted to each of the 66 species discussed, so that besides being interesting to read, it should be useful as a reference source for laymen. It is written in the easy style made popular in the author’s series The World of the Beaver, The World of the Red Fox, The World of the Raccoon, etc. Rue’s choice of mammals for this book is practical. He concentrates on large common species, including small mammals to round out the collection. If one is limited to one shrew, the short-tailed shrew Blarina brevicauda is the best-known example; if there are to be only two bats, the little brown (as a hibernating species) and the red (as a migrating one) are suitable representatives. It is odd, however, to have all marine mammals represented by Californian sea lions. Of the mammals discussed, only 13 are not found in Canada. The many illustrations in this book will make it especially appealing to children. The photographs, most of them excellent, are all by the author, who has obviously traveled widely in North America. The small maps of North America suffer the problems of BOTANY Plant Names By T.S. Lindsay. 1976. Facsimile of 1923 edition. Gale Research Co., Boston, Massachusetts. vii + 93 pp. $8. This small book, designed for gardeners and naturalists, deals with two topics: the principles of botanical nomenclature, and the origins and mean- ings of plant names. The International Code of Botanical Nomenclature did not exist when this book was first published. Some parts of Lindsay’s discussion of the rules of nomen- clature, based on the Vienna Rules of 1905, still apply, but other parts are no longer correct, e.g., “There is no accepted law as to names of varieties (1.e., cultivars). But botanists heed them not.” In 1923, the Primulales could correctly be designated by the now-obsolete term “cohort,” but even then the Primulaceae should not have been called a “division,” nor was the recognition of botanical varieties considered to violate the principle of binary nomenclature. Those all greatly reduced maps — the extent of the distribution of some species is underdone, and of others seemingly overdone. The badger and the cottontail are missing from Ontario, while the wolverine is represented in eastern Canada by a wide sweep of gray that belies its almost endangered status. The footprints are detailed but their arrangement into tracks is disappointing. Unless the gait an animal used to make the track is noted, it is usually impossible to tell which feet are which. For the black bear the forefeet are obviously different than the hind feet, but there is no legend indicating which is which, or how big either is. The appendices include a brief mention of what mammals may be seen in the many federal, state, and provincial parks on this continent; a compilation with addresses of the bureaus, departments, and agencies that deal with wildlife; and a list by family of all the mammals present in North America together with their scientific names. The reference list is short with the most recent item 1964, underlining that this printing of the 1967 edition has not been updated. ANNE INNIS DAGG Box 747, Waterloo, Ontario N2J 4C2 seeking an introduction to botanical nomenclature must look to more authoritatively written and more modern works. The names of plant genera, whether Latin or English, for which derivations are given are few, and appear to have been selected almost at random. Indexed under G, for example, there are only 21 Latin and 9 English names. Derivations of 10 of the Latin and 6 of the English names are scattered through the text; the others are merely classified as “commem- orative” or according to their linguistic origins. The chances are, therefore, that a generic or common name which one might wish to look up will not be included. Fortunately, several books that provide more extensive lists of meanings and derivations for botanical and common names are now available. Even a dictionary is a superior reference for this purpose. 334 The definitions of specific epithets are more numerous, although in this area, too, this book falls far short of more recent works. There are, moreover, many inaccurate or vague definitions, e.g., “oxy- canthus (sic), with sharp flowers”; “racemosus, full of clusters”; “repens, sudden, unexpected.” There are no illustrations. If no better works were available in 1923, this book Wildflowers across the Prairies By F. R. Vance, J. R. Jowsey, and J.S. McLean. 1977. Western Producer Prairie Books, Saskatoon. 214 pp., illus. Cloth $14.95; paper $8.95. In the words of the second author, “This book on wildflowers is, in general, a book to “open the eyes” of people of all ages as they look at the flowers of this region of North America. It is for travellers who come here from far lands, and for those who have lived on the prairies. It is a book for farmers and summer cottage residents, for those who visit our parks, and for those who walk or drive in cities and along the roads of Saskatchewan and neighboring areas. It is a book for all students of the natural world.” There is no doubt that the over 400 excellent color photographs will help serve this purpose in the prairies of Saskatchewan, Alberta, Manitoba, Montana, the Dakotas, and to a lesser extent beyond these limits. An informative text describes the plants and gives interesting notes on the habitats and areas where 186 species are found in Saskatchewan. Line drawings of flowers, seeds, or fruit accompany many of the species treated. These are of mixed quality, and indeed some might best have been omitted. Color photographs of related species are found throughout. These, although as good as those of the species being discussed, were Spring Flora of Wisconsin By Norman C. Fassett. 1976. 4th ed. (revised). Univer- sity of Wisconsin Press, Madison. 413 pp., illus. Paper $3.95; cloth $8.50. Previous editions of this little book which treats plants growing without cultivation and flowering before June 15th in Wisconsin, have proven to be immensely popular. This fourth edition is a greatly enlarged and revised version which has been prepared by Olive S. Thomson. In it, new species that represent recent introductions to the state have been added, descriptions have been enlarged, habitat and distri- bution data have been revised, over 100 new illustra- THE CANADIAN FIELD-NATURALIST Vol. 91 may then have been a useful addition to the popular literature of botany, but I see no reason to acquire it today. JAMES S. PRINGLE Royal Botanical Gardens, Box 399, Hamilton, Ontario L8N 3H8 found to be confusing by their presence on the page, even though they were mentioned in the text, and it seemed that they had been introduced because there was a picture available. It would have been better if these pictures had been given a page of their own, with their own descriptive text. Included in the volume are five pages of line drawings depicting various types of leaves, in- florescences, and flowers, which were reproduced from Wild Plants of the Canadian Prairies by A.C. Budd and K.F. Best. This is a most useful inclusion. Also to be found are a map depicting the area covered, a glossary, a short bibliography, an index to common and scientific names of species and families, and a color index which classifies the plants into four main flower color groups: red-pink, purple-blue, white, and greenish-yellow-cream. Fenton R. Vance took most of the photographs, but photo credits are given to some 30 additionai individuals. The line drawings were executed by J. S. McLean, and the text written by J. R. Jowsey. WILLIAM J. CODY Biosystematics Research Institute, Canada Agriculture, Ottawa, Ontario KIA 0C6 tions have been added, and some illustrations from previous editions have been redrawn. This most useful book will be welcomed in class and in the field by students not only in Wisconsin, but in the adjacent states and nearby parts of Canada. Naturalists in this region will also find this pocket- sized book a useful addition to their libraries even if they already possess an earlier edition. WILLIAM J. CODY Biosystematics Research Institute, Canada Department of Agriculture, Ottawa, Ontario KIA 0C6 1977 Canadian Forestry. The View beyond the Trees By C. R. Stanton. 1976. MacMillan, Toronto. 70 pp., illus. $5.95. The vital role of forests as national wealth is explained. Their management is of prime importance to derive benefits perennially. A historical back- ground is sketched beginning with the construction of houses in the past by Indian tribes. After the European settlements appeared, the shipbuilding industry grew up, utilizing timber. Later on lumber mills were established. Finally paper industry assumed major importance in national economy. A beautiful colored map of Canada is provided with forest regions and principal tree species, clearly marked. Typical pictures of each kind of vegetation were separately presented to give an idea how each region of vegetation looks like, e.g., boreal, montane, deciduous, and subalpine forest. Bargrams show the forest land in hectares and merchantable timber in cubic metres in each province of Canada. A short account appears on forest administration policy. As fire and pests cause maximum damage, management methods and intensive protection methods are required. New techniques of planting are being developed from time to time in order to keep pace with harvesting. Forest inventory work is speeded by ENVIRONMENT “Man of the Woods” By Herbert F. Keith. 1976. Syracuse University Press. Syracuse, New York. 164 pp. $5.95 (paperback edition of a 1972 publication) “Man of the Woods’ is a tale of what most people aspire to do with their lives, to be whom and do what they desire. Herbert Keith dreamed of being, and was, an Adirondack guide on the Oswegatchie River. His work is autobiographical of his life in and love for the town of Wanakena and the Oswegatchie River. His love for the region was kindled through his boyhood vacations to the lumber town of Wanakena, New York at its peak at the turn of this century. The early chapters of his work give an interesting account of Wanakena’s beginning and decline, which de- pended upon the production of the timber industry. It was this industry that first opened the region for tourism and guiding through the creation of railroad spurs to haul logs to the mills. It was during the timber and post-timber era that Keith dwells longest and introduces the reader to some of the more colorful BOOK REVIEWS 335 digitized stereoscope, which presents forest measure- ments from large-scale aerial photographs in computer-ready form. Mechanized logging opera- tions are carried out replacing the traditional lumberjack with his axe and saw. Veneers, plywoods, particle boards, pulp and paper, secondary wood, shingles, and maple syrup are some of the products of forests. At the same time, wildlife preservation is an adjunct function. Various research centres have been set up to tackle different problems. This book has numerous colored photographs of very high quality. It is not too technical; A layman could easily understand what forests are, how important they are for man’s survival, and their vital role in national economy. Any student (Grade 6 on) could get a bird’s-eye view of Canadian forests. I recommend that everyone take a glance at this title irrespective of their background, whether in biology, ecology, or forestry or whatever. We must con- gratulate C.R. Stanton for bringing out this concise edition. Dr. C. R. CHEVENDRA Science Librarian, University of Western Ontario, London, Canada characters and events of the time. His story-telling is based primarily on humor. There are moments of personal tragedy, however: the loss of a friend, the in-roads of an insatiable recreative society, and the loss of one man’s harmony with nature, which gives the reader a balanced view of the nature of wilderness living, a respect for all things great and small. The third period Keith dwells upon is that of the automobile. Keith dwelt longest on the first two periods perhaps because they were most memorable to him. Perhaps too, the age of the automobile gives him pain to have seen the careless casual destruction caused by overeager, overbearing, and undertrained modern man. Keith hones in on the problems associated with this new “sport” person in Chapters 18 and 19. I feel he tends to editorialize in these chapters and breaks out of his role as a story-teller, which disrupts the mood set throughout the rest of the work. Maybe this is Keith’s method to cause a self- 336 THE CANADIAN FIELD-NATURALIST evaluation of the reader’s attitude toward nature. Apart from Keith’s skills as a story-teller, the work contains a fine pictorial essay of the characters and way of life which compliments the story line at the moment. Another strong feature is the notes accompanying the work, referred to at the back of the text; these broaden its usefulness as a source of social and historical information. Also included are two maps. The first is of the railroad system on page 5, but it lacks a directional bearing and scale, which may Ecological Diversity By E. C. Pielou. 1975. Wiley, New York. 165 pp. $14.95. This book is primarily intended for studies of ecological communities. The questions confronted by an ecologist are (1) How many trophic levels are present, and are there a small number of intricately anastomosing food webs or a large number of simple unbranched food chains? (2) Do the species differ much in the amplitude of their tolerance ranges for various environmental variables? (3) Which of the species are autochthonous (evolved locally) and which allochthonous (evolved elsewhere) and which (if any) will soon become locally or globally extinct? (4) Are most or all of the community’s species fully adopted to the habit they occupy and to one another? The principles, explanations, applications, and derivations outlined in this book would throw some light on community studies. Moreover, the author clearly explains the difference between mathematical ecology and statistical ecology. The ecologists of the former category devise dynamic models suchas sets of differential-difference equations. Statistical ecologists on the other hand have less faith in conceptual models and the long chains of arguments arising from them. The author is of the opinion that the mathematicians run the risk of constructing interesting models divorced from reality while the statisticians run the risk of proving clear answers to ecologically un- interesting questions. I also fall in line with the firm conviction of the author that an ecologist with Vol. 91 limit its value. The second map on page 95 is a regional map which lacks a scale. Keith’s “Man in the Woods’ isa work well done and a fine piece of reading. It has something of interest for all. I highly recommend it. G. MADIGAN Box 39, MacDonald College, Woodland Resources, Ste. Anne de Bellevue, Quebec considerable field experience can recognize good questions and good answers. The book is divided into eight chapters opening with ‘Indices of diversity and evenness’ which covers Simpson Index, estimation of diversity of a large community, evenness and equitability, and hierar- chial diversity. Chapters two and three consisted of species-abundance distributions, followed by a testing hypothesis in chapter four. Spatial pattern, environ- mental gradients, local factors and global factors cover the other half of the book. An useful biblio- graphy is presented. Conventional subject and author indexes appear as usual. To me, it appears, a little background in mathematics is necessary to apply these principles because I (basically, I am a botanist) consulted a biometrician to explain some of the steps involved. This should not frighten an ecologist from using this book. A little guidance from a mathemati- cian will straighten your problems, if you have any. Taking into consideration all the above facts, this title is an excellent version on the topic in question. Diversity as a whole is treated in a coherent manner. It is highly desirable that a field ecologist be familiar with this type of knowledge. Undoubtedly, this is a piece of scholarly exposition by Ms. Pielou. C. R. CHEVENDRA Science Librarian, University of Western Ontario, London, Ontario Energy Flow — Its Biological Dimensions. A Summary of the IBP in Canada 1964-1974 Edited by T. W. M. Cameron and L. W. Billingsley. 1975. Royal Society of Canada, Ottawa. x + 319 pp. $5. The book consists of six sections corresponding to International Biological Program (IBP) outline: ter- restrial productivity, terrestrial conservation, fresh- water productivity, marine productivity, productivity processes, and human adaptability. The emphasis in all the 18 chapters is on the work carried out in Canada. The opening chapter is an overview pre- sented by W. H. Cook who had dealt with primary and secondary productivity, nitrogen fixation, terres- trial and aquatic environments, ecological sites and aT reserves, and human ecology. The first section constituted the studies on productivity, primary as well as secondary, and processes such as photo- synthesis and nitrogen fixation. The section on arctic ecosystem included the effects of population growth and the changing conditions on the Inuit hunting patterns, quantifies the present sources of food and economic conditions, and assesses their impact on wild life productivity. Specialized studies made in production processes, namely photosynthesis and nitrogen fixation, were dealt with in section two. The third section covered a national check-sheet survey of diverse ecological areas and indicated some represen- tative or unique sites recommended for preservation as ecological reserves. Legal and related requirements necessary for the effective preservation of such reserves were also presented. The fourth section is on freshwater productivity for which special environ- ments chosen are in a high arctic lake, and in a mountain lake in a more southerly latitude. Marine productivity was dealt with in section five and the projects included were from Gulf of St. Lawrence, Atlantic, Arctic, and Pacific coastal waters. All projects mostly concentrated on primary produc- tivity. The last section is on human adaptability. It dealt with Inuit population at Igloolik and formed part of an integrated international study of the circumpolar native peoples. This study had many aspects including demography, genetics, fitness, adap- BOOK REVIEWS 337 tations, and somatic growth patterns in the Inuit. Several independent studies were consolidated in order to compare growth patterns with different segments of the Canadian population. This book is a welcome contribution to the field of ecology and can be used as a textbook for any of the university courses. It is a fine collection of excellently written ecological contributions. The book is well edited and free of any printing errors and is recommended for research workers and postgraduate students. A complete schematic survey of IBP is presented. This isan example set forth by specialists in different fields of what we could achieve through international cooperation and standardization of methods and procedures. An integrated approach is well exemplified. It shows how scientific knowledge endeavors to improve the standard of living without exhausting the natural resources and how to maintain these perennially. Canadian participation in IBP and the results achieved are commendable. Some workers have not yet reported their projects’ progress. These may throw some light when they are consolidated and published sooner or later. No doubt we all have to appreciate the efforts of Canadian research teams involved in IBP. C. R. CHEVENDRA Science Librarian, University of Western Ontario, London, Ontario The Land that Never Melts: Auyuittuq National Park Edited by Roger Wilson. 1976. Peter Martin Associates with Publishing Centre, Supply and Services Canada. 212 pp. plus map, illus $5.95. Auyuittuq National Park on Baffin Island is dominated by glaciers flowing away from the central Penny Ice Cap, “powerful rivers of ice which have, over seemingly endless time, sculptured this pano- rama out of the hard rock of the peninsula.” This most northerly of Canadian parks illustrates many prin- ciples of geology, as Gifford Miller and Raymond Bradley explain in the first chapter. Erratics casually scattered in the park reflect their dispersal by glaciers, striations on bedrock indicate in what direction these glaciers moved, and slow-growing lichens indicate by their size how long the glaciers have been gone. One crustose lichen was 45 mm in diameter, indicating it had been growing on a glacier-free rock for 1200 years. The history of human occupation of the Arctic in general and of Baffin Island in particular is described in chapter 2 by Peter Schledermann who excavated a number of sites of Thule peoples, apparently the forebears of the Inuit who arrived from the west in the Baffin area about 1200 A.D. Such archeology does not yield the electrifying finds of earlier civilizations such as the Egyptian or the Mayan, but underlines the amazing feat of how man survived at all in such a hostile climate with no modern technology. Which of us could endure an arctic winter in a stone hut roofed with skin stretched over the bones of a whale we had caught with a home-made harpoon? Although all the many colored pictures in this book are superb, the photos of various habitats and arctic plants and the paintings of native birds by Jean-Luc Grondin in the third chapter “The Living Landscape” are even more breath-taking than usual. Patrick Baird discusses adaptations of plants that in general do not compete biologically with each other, as they do in southern Canada, but are rather pitted against their environment. Birds are easy to see and study because of the sparse vegetation, mammals less obvious except for the lemmings which every four years or so swarm 338 THE CANADIAN FIELD-NATURALIST in their myriads. The fourth chapter also by Patrick Baird gives helpful hints to park visitors — what to wear (thick clothing as a protection against cold and mosquitoes, and a lined jacket to cut the often gale-force winds); how to cross rivers with the least danger of becoming chilled; a caution not to camp on any vegetationless areas because here rockslides or snowslides probably occur. Plants and Animals of the Pacific Northwest By Eugene N. Kozloff. 1976. J. J. Douglass, Vancouver. 264 pp., illus. $17.50. The subtitle of this book reads “An Illustrated Guide to the Natural History of Western Oregon, Washington, and British Columbia.” In fact, the coverage extends from Vancouver, Nanaimo on Vancouver Island, south through Puget Sound and the Willamette Valley in Washington and Oregon. The intent of the book is to act as a reference and handbook for amateur naturalists and biology students. In this task, the book gets full marks. An earlier Kozloff book, entitled Seashore Life of Puget Sound, the Strait of Georgia and the San Jaun Archipelago, is a companion volume and a model for the present work. In both, Kozloff approaches natural history from the standpoint of examining habitats. Chapters consider the common plants and in- vertebrate animals of coniferous forests, open brushy places, wet places, backyards, vacant lots, and roadsides. A concluding chapter deals with verte- brate animals, but not thoroughly since other extensive treatments are readily available. There are over 320 color photographs and 125 line drawings, all of which are clear and leave no doubt as to the identity of the organism. The breadth of the book is impressive in terms of the number of taxonomic groups covered. Trees, shrubs, and common flowering plants are discussed, along with mosses, liverworts, ferns, and fungi. Common arthropods, snails, and slugs are also included. These are an unusual and welcome addition because these organisms are often discussed only in keys and academic tracts, largely inaccessible to Vol. 91 This pocket-sized book not only provides a superb field guide for those planning to travel in southern Baffin Island, but should encourage intrepid people to visit one of our newest and most exotic national parks. I recommend it highly. ANNE INNIS DAGG Box 747, Waterloo, Ontario N2J 4C2 amateur naturalists. The net result of Kozloffs work is a book that is very useful and comprehensive. Perhaps the most appealing aspect of the book is Kozloff’s fresh, personal, and interesting writing style. Discussions of plants and animals come to the reader as though presented by a warm and wise old mentor. This, for the amateur, is far more appealing than dry descriptive text. Added to this is the handsome presentation of the book. One of the problems, aside from the price, is that the book has to be read right through to be appreciated or truly useful. There are no keys, and plants and animals are not discussed as groups. If an unknown flowering plant is found, for example, the reader may have to check each chapter to find its identity. This is partially circumvented by discussions of habitats, but un- trained observers cannot really be expected to be highly discriminatory in this respect. Also, the book’s bulk may present a problem for field use. Kozloffs previous book is already a standard reference for seashore life, and his Plants and Animals may become one as well, if it is reprinted in good paperback and reasonably priced. It will always have limited usefulness to Canadians on the west coast, especially if they live or travel farther north than the Vancouver area, but for now, it is one of the few books that offer a comprehensive natural history on the central west coast. BRIAN D. WILKES 688A Winchester Avenue, Nanaimo, British Columbia VOR 4B8 LOW NEW TITLES Zoology Adaptations to environment. Essays on the physiology of marine animals. 1976. Edited by R. C. Newell. Butter- worths, Boston. xv + 264 pp., illus. Paper $10.95. {The African buffalo. A study of resource limitation of populations. 1977. By A.R.E. Sinclair. University of Chicago Press, Chicago. xii + 355 pp., illus. $17.50. *Alberta birds 1961—1970. 1976. By T.S. Sadler and M.T. Myres. Occasional Paper Number |. Provincial Museum of Alberta, Edmonton. $3.25. *An American crusade for wildlife. 1975. By James B. Toefether. Wincester Press (Canadian distributor Nelson, Foster and Scott Willowdale). xii + 409 pp., illus. $14.50. {Animal communication. 1977. By Hubert and Mable Frings. 2nd edition. University of Oklahoma Press, Norman (Canadian distributor Burns and MacEachern, Don Mills). xi + 207 pp., illus. Paper $6.25. The Audubon wildlife treasury. 1976. Edited by L. Line. Lippincott, New York. 192 pp. $15.95. The bee book. The history and natural history of the honey- bee. 1976. By Daphne More. University Books, New York. 144 pp., illus. $12.50. {Biological insect pest suppression. 1977. By H. C. Coppel and J.W. Mertins. Advanced Series in Agricultural Sciences 4. Springer-Verlag, New York. xiii + 314 pp., illus. $29.60. *Biology of insects. 1976. By David J. Horn. Saunders, Philadelphia. vii + 439 pp. $14.50. The bird table book in colour. 1977. By Tony Soper. Illus- trated by R. Gillmor. David and Charles, Newton Abbot, England. 128 pp. £2.95. Birds as builders. 1977. By Peter Goodfellow. David and Charles, Newton Abbot, England. 160 pp., illus. £4.95. British opisthobranch molluscs. Mollusca — Gastropoda. Keys and notes for the identification of species. 1976. By T. E. Thompson and G. H. Brown. Academic, New York. vill + 204 pp., illus. Paper $7.65. Crows, jays, ravens and their relatives. 1977. By Sylvia Bruce Wilmore. David and Charles, Newton Abbot, Eng- land. 192 pp., illus. £5.95. {The earthworms (Lumbricidae and Sparganophilidae) of Ontario. 1977. By John W. Reynolds. Illustrated by Daniel L. Dindal. Life Sciences Miscellaneous Publications. Royal Ontario Museum, Toronto. ix + 141 pp. $8. BOOK REVIEWS 339 Endangered and threatened amphibians and reptiles in the United States. 1976. Compiled by R. E. Ashton. Edited by S. R. Edwards and G. R. Pisani. Herpetological Circular Number 5. Society for the Study of Amphibians and Rep- tiles, Lawrence, Kansas. 65 pp. +Energetic significance of the annelids and arthropods in a Swedish grassland soil. 1977. By T. Persson and U. Lohm. Ecological Bulletins NFR 23. Swedish Natural Science Research Council, Stockholm. 211 pp. The ethology of predation. 1976. By E. Curio. Springer- Verlag, Berlin. x + 250 pp. $29.60. Fish population dynamics. 1977. Edited by J. A. Gulland. Wiley, New York. xi + 372 pp. $25. Fossil animal remains. Their preparation and conservation. 1976. By A. E. Rixon. Athlone, London viii + 304 pp., illus. Paper $12. +A guide to bird finding east of the Mississippi. 1977. By Olin Sewall Pettingill, Jr. Illustrated by G. M. Sutton. 2nd edition. Oxford University Press, New York. xvii + 689 pp. $15.95. A guide to the fishes of the temperate Atlantic coast. 1977. By Michael J. Ursin. Clarke Irwin, Toronto. Cloth $10.75; paper $6.75. t+Guide to the pigeons of the world. 1976. By Andrew MeNeillie. Elsevier (Canadian distributor Burns and Mac- Eachern, Don Mills). 160 pp., illus. Paper $5.95. Land hermit crabs. 1976. By Paul J. Nash. Clarke Irwin, Toronto. $1.95. tLarvae of the North American caddisfly genera (Tri- choptera). 1977. By Glenn B. Wiggins. Illustrated by Anker Odum. University of Toronto Press, Toronto. xi+ 401 pp. $25. Legion of the night: the underwing moths. 1976. By T. D. Sargent. University of Massachusetts Press, Amherst. xill + 222 pp., illus. $15. The mayflies of North and Central America. 1976. By G. F. Edmunds, Jr., S. L. Jensen, and L. Berner. University of Minnesota Press, Minneapolis. x + 330 pp., illus. $28.50. {Minnesota birds. Where, when, and how many. 1975. By Janet C. Green and Robert B. Janssen. University of Minnesota Press, Minneapolis (Canadian distributor Burns and MacEachern, Don Mills). 217 pp., illus. $9.75. Moments of discovery: adventures with American birds. 1976. By E. Ponter and M. Harwood. Dutton, New York. $29.95. 340 A natural history of marine mammals. 1976. By Victor B. Scheffer. Illustrations by Peter Parnall. Scribner, New York. xu + 158 pp. $7.95. tNorth American bird songs — a world of music. 1977. By Poul Bodesen. Scandinavian Science Press. Klampen- borg, Denmark. 254 pp., illus. $15.50. Nudibranchs. 1976. By T. E. Thompson. Clarke Irwin, Toronto. Paper $5.95. Ontario Nest Records Scheme. 1977. By George K. Peck. Thirteenth Report (1956 — 1976). Royal Ontario Museum, Toronto and Canadian Wildlife Service, Ottawa. 34 pp. { Optical signals. Animal communication and light. 1977. By Jack P. Hailman. Indiana University Press, Bloomington (Canadian distributor Fitzhenry and Whiteside, Pickering, Ontario). xix + 362 pp., illus. $15. {The Passenger Pigeon. Its natural history and extinction. 1955, 1973. By A. W. Schorger. University of Oklahoma Press, Norman. (Canadian distributor Burns and Mac- Eachern, Don Mills). xiii + 424 pp., illus. Paper $7.50. {Précis de zoologie: vertébrés. 2. Reproduction, biologie, évolution et systématique. Agnathes, poissons, amphibiens et reptiles. 1976. Par P.-P. Grassé. 2e édition. Masson, Paris. 480 pp., illus. +Précis de zoologie: vertébrés. 3. Reproduction, biologie, évolution et systématique. Oiseaux et mammiféres. 1977. Par P.-P. Grassé. 2e édition. Masson, Paris. 408 pp., illus. 130F. tA second book of Canadian mammals. 1977. By Charles Paul May. Illustrated by John Crosby. 2nd edition. MacMillan, Toronto. 109 pp. Paper $4.95. Sex lives of animals without backbones. 1976. By H. H. Najarian. Scribner, New York. x + 116 pp., illus. $6.95. Survey of the world’s aphids. 1976. By V.F. Eastop and D. H. Rislanders. Junk, The Hague, vi + 574 pp. Dfl 160. There’s a raccoon in my parka. Adventures with wildlife in British Columbia, Alaska and the Yukon. 1977. By Lyn Hancock. Doubleday, Toronto. $8.95. *Waterfowl studies. For the decoy maker and collector, hunter, bird watcher, waterfowl artist and naturalist. 1976. By Bruce Burk. Wincester (Canadian distributor Nelson, Foster and Scott, Willowdale). xiv + 254 pp., illus. $17.95. +Ways of the six-footed. A delightful introduction to the world of insects. 1977. By A. B. Comstock. Corness Uni- versity Press, Ithaca. xviii + 152 pp., illus. $5.95. The web of adaptation: bird studies in the American tropics. 1976. Quadrangle, New York. 176 pp. $8.95. THE CANADIAN FIELD-NATURALIST Vol. 91 *Wildlife management in Europe. 1977. By Anne Innis Dagg. Otter Press, Waterloo. ix + 324 pp. Paper $6.50 + 50¢ postage. Botany Atlas of United States trees. Volume 3. Minor western hardwoods. 1976. By E. L. Little. Miscellaneous Publica- tion Number 1314. United States Department of Agri- culture, Washington. 13 pp. + 290 maps. $9.10. {The Enterprise, Wisconsin radiation forest. Radioecology studies, part 2. 1977. Edited by J. Zavitkovski. United States Technical Information Center, Oak Ridge, Ten- nesee. 220 pp., illus. $7.50. Green worlds: plants and forest life. 1976. By D. Bellamy and M. Boorer. Doubleday, Garden City, New York. 144 pp. $14.95. Index to plant distribution maps in North American periodicals through 1972. 1976. Compiled by W.L. Phillips and R. L. Stuckey. Hall, Boston. 686 pp. Intermountain flora. Vascular plants of the intermountain west, U.S.A. Volume 6. Monocotyledons. 1977. By A. Cronquist, A. H. Holmgren, N. H. Holmgren, J. L. Reveal and P.K. Holmgren. Columbia University Press, New York. 584 pp. $35. Mount Revelstoke National Park wild flowers. 1976. By James H. Soper and Adam F. Szazawinshi. Natural History Series Number 3. National Museums of Canada, Ottawa. 96 pp., illus. $2.50. *Seeds and fruits of plants of eastern Canada and north- eastern United States. 1977. By F.H. Montgomery. University of Toronto Press, Toronto. xi + 232 pp., illus. $25. Trees and shrubs of the United States. A bibliography for identification. 1976. By E. L. Little and B. H. Honkala. Miscellaneous Publication Number 1336. United States Department of Agriculture, Washington. 55 pp. $1. Tropical trees: variation, breeding and conservation. 1976. Edited by J. Burley and B. T. Styles. Linnean Society Symposium Series Number 2, Proceedings of an Inter- national Symposium, Oxford University, England, April, 1975. Academic, New York. xviii + 244 pp. $20.25. tVascular plants of British Columbia. A descriptive resource inventory. 1977. By Roy L. Taylor and Bruce MacBryde. Technical Bulletin Number 4. University of British Columbia Press, Vancouver. xxiv + 754 pp. Paper $28. Vascular plants of the Nevada test site and central- south Nevada: ecological and geographic distributions. 1976. By J. C. Beatley. United States National Technical Information Service, Springfield, Virginia. 316 pp. $12.25. NOT The vascular plants of South Dakota. 1976. By T. Van Bruggen. Iowa State University Press, Ames. xxvi + 538 pp. Paper $7.95. *Wildflowers across the prairies. 1977. By Fenton R. Vance, James R. Jowsey and James S. McLean. Western Producer Prairie Books, Saskatoon. 214 pp., illus. Cloth $14.95; paper $8.95. Wildflowers of the southeastern United States. 1975. By W.H. Duncan and L. E. Foot. University of Georgia Press, Athens. vii + 296 pp. $12. Environment {Canada’s threatened species and habitats. 1977. Edited by Theodore Mosquin and Cecile Suchal. Proceedings of the First National Conference on Canada’s Threatened Species and Habitats, Ottawa, May, 1976. Canadian Nature Federation. Ottawa.-x + 185 pp. $8. *Erosion of land in northwestern Alberta. 1976. By the Alberta Environmental Conservation Authority, Ed- monton. 73 pp. Paper free. Life in the future: prospects for man and nature. 1976. By M. Ross-Macdonald, M. Hassell, and S. McNeill. Double- day, Garden City, New York. 64 pp. $14.95. Managing the environment. Towards a comprehensive approach. 1976. By P. W. House. Teakfield, Farnborough, England. *Mankind’s future in the Pacific. 1977. Edited by R. F. Scagel. University of British Columbia Press, Vancouver. The plenary and special lectures of the 13th Pacific Science Conference, August, 1975. ix +206 pp. Paper $6.95. {Mathematical ecology. 1976. By E. C. Pielou. Wiley, New York. x + 385 pp. $19.50. Nature in trust: the history of nature conservation in Britain. 1976. By John Sheail. Blackie, Glasgow. xiv + 270 pp., illus. £5.95. Our continent. A natural history of North America. 1976. By the National Geographic Society, Washington. 398 pp., illus. $11.95. Ponds. Their wildlife and upkeep. 1977. By R. Burton. David and Charles, Newton Abbot, England. 176 pp., illus. £4.50. *Structure and function of tundra ecosystems. 1975. Edited by T. Rosswall and O. W. Heal. Papers presented at the IBP Tundra Biome V International Meeting on Bio- logical Productivity of Tundra, Abisko, Sweden, April, 1974. Ecological Bulletins NFR 20. Swedish Natural BOOK REVIEWS 34] Science Research Council, Stockholm. 455 pp. 50 Skr. Texas wild: the land, plants and animals of the lone star state. 1976. By R. Phelan and J. Bones. Dutton, New York. $30. The web of life: the ecology of earth. 1976. By J. D. Oates, D. Toomer, and A. Cane. Doubleday, Garden City, New York. 144 pp. $14.95. Miscellaneous {The backpacker. 1972, 1977. By Albert Saijo. Van Nostrand Reinhold, Toronto. 192 pp. Paper $3.50. {Cross-country Canada. Handbook and trail guide for cross-country skiers. 1977. Van Nostrand Reinhold, Toronto. xii + 212 pp., illus. Paper $6.95. Discover the sky with telescope and camera. 1976. By R. Knox. Clarke Irwin, Toronto. $9.95. Early wildlife photographers. 1977. By C. A. W. Guggis- berg. David and Charles, Newton Abbot, England. 128 pp., illus. £4.50. Evolution and the diversity of life. Selected essays. 1976. By Ernest Mayr. Belknap Press of Harvard University Press, Cambridge, Massachusetts. xii + 722 pp., illus. $20. + Geographic variation, speciation and clines. 1977. By John A. Endler. Princeton University Press, Princeton. 246 pp., illus. Cloth $16; paper $6.95. Introduction to natural selection. 1976. By C. Johnson. University Park Press, Baltimore. x + 214 pp. $12.50. tIssues in outdoor recreation. 1977. Edited by C. R. Jensen and C. T. Thorstenson. 2nd edition. Burgess, Minneapolis. ix + 326 pp. Paper $7.95. {The mineralogy of Arizona. 1977. By J. W. Anthony, S. A. Williams, and R. A. Bideau. University of Arizona Press, Tucson. 241 pp. + maps. Cloth $22.50; paper $9.75. tOutdoor recreation in America. Trends, problems and opportunities. 1977. By C. R. Jensen. 3rd edition. Burgess, Minneapolis. viii + 269 pp., illus. $12.95. The science of speleology. 1976. Edited by T. D. Ford and C. H. D. Cullingford. Academic, New York. xiv + 594 pp., illus. $29.50. {Theories of populations in biological communities. 1977. By T. M. Fenchel. Ecological Studies Volume 20. Springer- Verlag, New York. x + 144 pp. $27.30. tavailable for review *assigned for review Instructions to Contributors Content The Canadian Field- Naturalist is a medium for publica- tion of original scientific research papers in all fields of natural history that have relevance to Canada. 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The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain the scientific quality and overall high standards of the journal. TABLE OF CONTENTS (Concluded) Changes in the avifauna of the West Foxe Islands, Northwest Territories, 1956-1976 F. G. COOCH Unusual predators of Snow Goose eggs KENNETH F. ABRAHAM, PIERRE MINEAU, and FRED COOKE New and notable finds in the Alaskan vascular flora G. HALLIDAY Range extensions of the Water Shrew and Mink Frog in the James Bay region of Quebec D. J. OXLEY, R. A. CouTTs, and N. G. H. BOYLE Range of the Bushy-tailed Wood Rat (Neotoma cinerea) in Alberta DAVID W. KRAUSE and BRUCE G. NAYLOR News and Comment Book Reviews Zoology: Biology of Insects — North American game birds of upland and shoreline — Waterfowl studies — An American crusade for wildlife — America’s master of bee culture. The life of L. L. Langstroth — Biology of the Kaminuriak population of barren-ground caribou. Part 4: growth, reproduction and energy reserves — Répartition géographique des poissons, leur abondance relative et bathymétrie de la région du Lac Saint-Pierre — Cartes de répartition géographique des espéces de poissons au sud du Québec, d’apres les inventaires ichthyologiques effectués de 1963 a 1972 — Les poissons de la région de Montréal, la péche sportive et commerciale, les ensemencements, les frayeres, la contamination par le mercure et les PCB — Méthodes de péche expérimentale, en eau douce, a l’usage du biologiste et du technicien de la faune — The anatomy of fishes. Parts I and 1] — Wolf... Kill! The wilderness called Shunka — A field guide to birds’ nests — Pictorial guide to the mammals of North America. Botany: Plant names — Wildflowers across the prairies — Spring flora of Wisconsin — Canadian forestry. The view beyond the trees. Environment — “Man of the Woods” — Ecological diversity — Energy flow - its biological dimen- sions. A summary of the IBP in Canada 1964-1974 — The land that never melts: Auyuittuq National Park — Plants and animals of the Pacific Northwest. New Titles Mailing date of previous issue 28 June 1977 314 StF BS 322 323 325 326 THE CANADIAN FIELD-NATURALIST Volume 91, Number 3 1977 Editorials Reporting of range extensions DAVID P. SCOTT BANS) Editor’s comments and policy LORRAINE C. SMITH 221 Articles Shorebirds at Long Point, Lake Erie, 1966-1971: seasonal occurrence, habitat preference, and variation in abundance MICHAEL S. W. BRADSTREET, GARY W. PAGE, and W. GAVIN JOHNSTON DDS Population dynamics, home ranges, and habitat associations of the Yellow-cheeked Vole, Microtus xanthognathus, in the Northwest Territories RICHARD J. DOUGLASS 23), Nesting and brood ecology of Lesser Scaup at Waterhen Marsh, Saskatchewan JAMES E. HINES 248 The genus Crangonyx (Amphipoda: Gammaridae) in the central Connecticut River system DOUGLAS G. SMITH 256 Pteridophytes of the Regional Municipality of Waterloo, Ontario CRAIG A. CAMPBELL and DONALD M. BRITTON 262 The Biological Flora of Canada — A new series GEORGE H. LA ROI 269 Movements and habitat use among interacting Peromyscus leucopus as revealed by radiotelemetry DALE M. MADISON 273 Prairie fires and pronghorn use of cactus JOHN G. STELFOX and HAROLD G. VRIEND 282 Notes Waterfowl use of exotic wild rice habitat in northern Saskatchewan DONALD G. PEDEN 286 The Wolffish, cf. Anarhichas denticulatus, new to the Amundsen Gulf area, Northwest Territories, and a probable prey of the Ringed Seal THOMAS G. SMITH 288 Locations of winter dens utilized by striped skunks in Delta Marsh, Manitoba GRAHAM R. P. MUTCH 289 Age and fecundity of the Tadpole Madtom, Noturus gyrinus, on Long Point, Lake Erie : ROBIN MAHON 292 The Little Gull (Larus minutus) in Arctic North America STEPHEN R. JOHNSON and WILLIAM J. ADAMS 294 Summer food habits of Golden Eagles in southwestern Alberta D. A. BOAG 296 The occurrence of a narwhal(Monodon monoceros) in Prince Albert Sound, western Victoria Island, Northwest Territories THOMAS G. SMITH 299 Distribution of stream salamanders in southwestern Quebec WAYNE F. WELLER 299 The river otter (Lutra canadensis) on the north slope of the Brooks Range, Alaska AUDREY M. MAGOUN and PATRICK VALKENBURG 303 Lichens of the Bamfield Marine Station, Vancouver Island, British Columbia FRAN BENTON, IRWIN M. BRODO, and DAVID H. S. RICHARDSON 305 Narwhals (Monodon monoceros) observed near King Christian Island, Northwest Territories NICHOLAS A. ROE and WILLIAM J. STEPHEN 309 A Zone-tailed Hawk in Nova Scotia IAN A. MCLAREN and ANDREW MACINNES 310 Summer use of a highway crossing by mountain caribou DONALD R. JOHNSON and MICHAEL C. TODD 312 ISSN 0008-3550 concluded on inside back cover the CANADIAN ~ FIELD-NATURALIST | MUS, COMP. Z Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada= TK Volume 91, Number 4 October-December 1977 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Their Excellencies the Governor General and Madame Jules Léger The objectives of this Club shall be to promote the appreciation, preservation, and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and co-operate with organizations engaged in preserving, maintaining, or restoring environments of high quality for living things. Members of Council* President: R. A. Foxall E. Beaubien J. Murray W. J. Cod M.N Vice President: R. Taylor j He G Neuen Recording Secretary: D. R. Laubitz A. Dugal G. Oyen : A. J. Erskine G. Patenaude : S. Armst Corresponding Secretary S. Armstrong ©, Gada 1K scans Treasurer: P. J. Sims J. E. Harrison S. M. Teeple Past President: E. C. D. Todd B. Henson C. G. van Zyll de Jong H. N. MacKenzie Correspondence: Address to The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada KI1Y 4J5 The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club with the assistance of contributions from the National Research Council of Canada and The Canadian National Sportsmen’s Show. Opinions and ideas expressed in this journal are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Editor: Lorraine C. Smith Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy Associate Editors C. D. Bird A. J. Erskine David P. Scott E. L. Bousfield Charles Jonkel Stephen M. Smith Francis R. Cook Charles J. Krebs Robert E. Wrigley George H. La Roi Copy Editor: Marilyn D. Dadswell Business Manager: W. J. Cody Production Manager: Pauline A. Smith Box 3264, Postal Station C Chairman, Publications Committee: C. G. van Zyll de Jong Ottawa, Canada K1Y 4J5 Subscriptions and Membership Subscription rates for individuals are $7.00 per calendar year. Libraries and other institutions may subscribe at the rate of $15.00 per year (volume). The annual membership fee of $7.00 includes club publications. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be‘mailed to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada K1Y 4J5. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Back Numbers Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field-Naturalists’ Club, 1879-1886, and The Ottawa Naturalist, 1887-1919, may be purchased from the Business Manager. Address manuscripts on birds to the Associate Editor for Ornithology: Dr. A. J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick EOA 3C0 All other material intended for publication should be addressed to the Editor: Dr. Lorraine C. Smith, R. R. 3, Stittsville, Ontario, Canada KOA 3G0 Cover: White-tailed Ptarmigan photographed by Donald A. Smith in the Rocky Mountains on 20 June 1968. See article on page 367. The Canadian Field-Naturalist Volume 91, Number 4 October-December 1977 The Myth of the Non-Consumptive User BRIAN WILKES Nanaimo Naturalist Club, Box 125, Nanaimo, British Columbia V9R 5K4 Wilkes, Brian. 1977. The myth of the non-consumptive user. Canadian Field-Naturalist 91(4): 343-349. The concept that some outdoor recreational activities are non-consumptive of the resource base is examined and rejected. Typical non-consumptive activities are seen to be consumptive along spatial, temporal, and physical dimensions. The wide acceptance of this erroneous concept has led to inappropriate behaviors on the part of Naturalist Club and Federation members. Serious errors in park and natural area planning and management have been made. Rejection of the concept frees us to formulate new guidelines and planning tools for parks and similar reserves. User restrictions, a proposed theory of non-use planning, and a new justification for landscape preservation are discussed within this context. Key Words: non-consumptive user, recreation, resource base, parks, natural areas, landscape preservation. Each year, hundreds of thousands of people participate in various outdoor recreational acti- vities, but neither hunt nor fish, and are generally referred to as “non-consumptive” users. The purpose of this article is to cast serious doubt on the validity of this term. The per- spective presented here is a policy statement adopted in principle by the Vancouver Island Region of the Federation of British Columbia Naturalists. The article was written at the suggestion of Neil Dawe, the past Regional Vice- President, and was subsequently reviewed and edited by a committee of executive officers. In this article, consumption is discussed only in terms of outdoor recreation usually found in natural areas, parks, reserves, or conservation areas, and in unprotected semi-wilderness. Con- sumption in terms of what it takes to maintain the essentials of human life, such as food, water, or oxygen is not considered. The points pre- sented here have important implications for naturalist clubs and conservation groups across Canada. It is not new to question the idea of the non-consumptive user, but there seems to have been no national discussion on the issue. Hopefully, this article will generate one. It is easy to understand why recreational hunting and fishing are considered consumptive. Living organisms are physically removed from the scene, and consequences are apparent when populations of game decline. These resources are supposedly renewable, and can be manipulated by some sort of conventional management. Certain conservation groups, or individuals in them, often rail against consumptive forms of recreation. Naturalist groups typically cast themselves in this light. Other groups recognize and accept the consumptive nature of their activity, arguing that they merely crop off some sort of “harvestable surplus.” Rod and gun clubs, rifle associations and other groups are in this category. No matter what one’s particular attitude is toward hunting and fishing, there is general agreement that these are consumptive activities. They are closely regulated in terms of bag limits or in the number of licensed parti- cipants. These controls derive from the recogni- tion of the consumptive nature of the activity, and are consistent with conventional manage- ment techniques. By contrast, hiking or back-packing, sight- seeing, general tourism and camping in parks, 343 344 nature study, nature photography, and pic- nicking are clearly regarded as non-consumptive of the resource base. These particular activities, and all the others in this category, do not seem to remove living organisms from the scene. They are regarded as healthful pursuits that are benign in terms of the surrounding landscape. Parti- cipants in these activities are regarded as non- consumptive users of outdoor recreation re- sources, and consequently there are few controls governing their numbers or behavior. The non-consumptive user can be categorized in a number of different ways according to the frequency and duration of participation in conventional non-consumptive activities. Natu- ralist clubs and hiking clubs which organize a specific roster of regular outings are one such category; others include the cubs, scouts, and girl guides, summer camps, wilderness users, recrea- tion vehicle enthusiasts, etc. By far the largest category consists of all the people who camp in or otherwise visit national, provincial, or regional parks, or who, in British Columbia and other provinces, travel on logging roads and camp along the way. It is difficult to say precisely what the size of this group is. No one can possibly guess the total number of people who visit unorganized faci- lities, or who seek out other sorts of crown wildland for purposes of recreation. The point is that non-consumptive users are present in far greater numbers than consumptive users. For example, in 1975, the number of hunters and anglers in British Columbia was about 512 000. No figures are available on the numbers of hunter or angler days for that year, but they could not possibly aproach the 8.7 million day and overnight visits to provincial parks in British Columbia for the same year.! In addition, the British Columbia Forest Service provides un- supervised camping facilities throughout the province, and cannot estimate the number of people who use them. Both Crown Zellerbach and MacMillan Bloedel provide limited facilities in their timber limits, but lost count of the number of users when access to major logging ‘Figures for hunters and anglers provided by G. Reid, British Columbia Fish and Wildlife Branch, Nanaimo. Figures on park use supplied by Mick Collins, Research Section, British Columbia Parks Branch. THE CANADIAN FIELD-NATURALIST Vol. 91 roads opened on a twenty-four hour basis a few years ago. They do estimate, however, the yearly visitation to be in the tens of thousands. ; These are estimates for British Columbia alone, with a population of 2.5 million. We would surmise that the total number of “non- consumers” ranging across the landscape of all provinces and territories is staggering — far greater than park visitor statistics and parti- cipation rate data from surveys suggest. Entire industries and retail empires have been created to cater to the needs of back-packers and canoe trippers alone. The “non-consumers” are using more sophisticated and mechanized equipment every year. It is now necessary for naturalist and conservation groups to recognize that non- consumptive users are no longer a few groups of nature buffs or boy scouts hiking into the hills. They are present in this country in enormous numbers. They have become big business, and a big problem. Is the notion of non-consumptive use valid? The very idea of it does not seem to square well with recognition that some recreational land- scapes become seriously degraded over a period of time. But strangely enough, the idea has become firmly imbedded in the rhetoric of the conservation movement. Most conservation of- ficers and professional biologists use the term to refer to groups of non-hunters and non-anglers. Even our own Federation referred to its members as non-consumptive users in a recent brief on the Fish and Wildlife Branch to the Minister of Conservation and Recreation. Per- haps it is time that we, as naturalists, escaped from this comfortable illusion. The concept of the non-consumptive user of outdoor recreation resources is false. It is an outdated concept that appears bankrupt of accuracy. Much like the now discredited notion of multiple use, the concept may have been useful once, but now in the face of the information available, it must be discarded. In fact, because the notion of the non-consumptive user has been so widely accepted, many serious errors have been made in land-use planning and in the philosophy of the conservation movement. Here is a case in which a comfortable myth has been applied as a principle of land use, and as a result some of the major objectives of the conservation movement are in jeopardy. 1977 Natural history clubs and federations have a major voice in the effective criticism of indus- trial, institutional, and private resource con- sumption issues. Until now, members of these organizations could lob their criticism from the safe fortress of the notion that their own activities were non-consumptive. But if non- consumption is a myth, then we can be held up to ridicule by our opponents.? If so-called non-consumptive activities are not so benign, then we had better acknowledge this and get down to the serious business of re- assessing our priorities. One of the major objectives of the conservation movement, and one behind which naturalists stand firmly, is the preservation of natural landscapes and habitats. We have focused on gaining legislative protec- tion for them without very seriously addressing the question of what happens to them next. We have not only supported the preservation of these lands, we have also been guilty of en- couraging their “non-consumptive” use by our own members and the general public. In fact, the chief argument used in support of natural area preservation, except ecological reserves, is the benefit that supposedly accrues to the public in terms of recreation. This argument will have to be abandoned, particularly if clubs and federa- tions address the contradiction of supporting both the preservation and use of natural land- scapes. Non-consumptive users do consume recrea- tion resources along spatial, visual, and physical dimensions. They trample and _ re-arrange vegetation patterns, disturb wildlife, and are the chief distributors of refuse across the land. Let us discuss these and then consider some important implications of the position taken. *The Village Lake Louise conflict is a good example. That was one of the major conservation victories in Canada. The environmental grounds against the proposed ski resort development were framed in terms of lost scenic and ecological amenities to non-consumptive users. The fact that it was ina national park only gave weight to our arguments. But if the project proponents, Imperial Oil et al., could have identified the long-term damage by tourists and scenery gawkers to the amenities of the site, and shown that this damage would be almost guaranteed by the management philosophy of Parks Canada, then the outcome of the conflict may have been very different. See Nature Canada 1(11): 35 and 1(2): 33, 1972. WILKES: MYTH OF NON-CONSUMPTIVE USER 345 Spatial consumption simply means recreation consumes space. Picture a natural landscape. In order for it to be of any conventional recrea- tional use, arrangements must be made for access to it and probably for accommodation in it. This results in the direct physical consumption of habitat in the area. A small park might serve as an illustration of this. Ivy Green Provincial Park, south of Nanaimo on Vancouver Island, is sixty- two acres in size and is bisected by the Island Highway. The park is classified as Class A or dedicated to the preservation of the natural environment (per the Park Act for the Province). Ivy Green contains forty-eight campsites, each about 108 m? (1200 ft?), thirty toilets and associated facilities located on pads cleared in the forest, a trailer sani-station, a paved parking lot for 104 cars, 3.5 km (2.2 mi) of roads with about 7.5-m (25-ft) clearances, and a large service yard (from the British Columbia Parks Data Handbook). All these facilities are installed for the non-consumptive user, and the process of installing them has left only about a quarter of the park unimpaired — and this is squeezed in between the campsites. The act of providing for the accommodation of non-consumptive users has succeeded in directly consuming three-quarters of the habitat in a park which has statutory protection from impairment. In this example, the visitors do not directly remove organisms or entire habitats from the scene. The government does it for them, with our blessings. How many small parks can you think of in similar circumstances? The problem of spatial consumption also extends to mechanized recreational vehicles. Trail bikers, waterskiers, and snowmobilers require much more space than do hikers or canoeists. Mechanized and non-mechanized pursuits rarely mix well on the same land- scape at the same time because of conflicts between users. Therefore, most areas are needed to accommodate all the participants of all the various activities, and when more space is needed, it means that what was formerly available has been used up. Conflicting acti- vities might be regulated in the same space by requiring that each occur at different times of the day or week. But this turns into a problem of regulations and controls, which appear to be unacceptable to those recreating under the 346 illusion that their own activity is not con- sumptive at all. Visual consumption means that large numbers of people consume solitude. Recreation research: has confirmed that human crowding in outdoor recreation settings results in decreased satisfac- tion with the experience in the mind of each visitor (Lucas 1964; Stone and Taves 1956). Crowds in any particular area can build to the point where the scenic amenities of the site are completely lost by the presence of too many people. The visual and auditory impact of crowding results in a general feeling of dissatis- faction with the setting, and is often referred to as perceptual carrying capacity (Bouchard 1973; Lime and Stankey 1971). The problem is that large numbers of people (or even small numbers, depending on your degree of “purism”) make solitude scarce. Since the necessary precondition of scarcity is consumption, we can conclude that visual resources have been consumed when the scenic amenties of a site are lost. If solitude becomes scarce enough to engender a feeling of over-crowding amongst people at a particular site, perceived or “expected” space has been all used up. As often as not, it is solitude that people seek when they visit natural landscapes. Here isa case in which larger numbers consume the very quality sought. Another aspect of visual consumption is the visual impact humans have on wildlife. There are a number of wildlife species that seem to require privacy from human intrusion in order to thrive in their respective ecosystems. In these cases the presence of people may not be directly con- sumptive, but in the long run the result is the same. These organisms do not have somewhere else to go when they are pushed out by human presence. They are where they are because that is where they must be. The alternative for them is simply to disappear. Beyond requirements for access and accom- modation on recreational landscapes lies the problem of direct physical impact. This was first identified as a problem as early as 1929 (Bates 1935; Meinecke 1929). Subsequent studies have shown that in certain environments, such as forested areas with a well developed ground cover, very severe impacts occur with the lightest use, and that physical impact is cumulative over a period of time (Frissell and Duncan 1965; La THE CANADIAN FIELD-NATURALIST Vol. 91 Page 1967). Cumulative impact simply means that the year-to-year effect of human presence in natural recreation settings continues to build gradually until serious changes become ap- parent. Even light and occasional use of an area for hiking or nature study can have its effect in time. The amount of effect is also dependent on the particular sensitivity of the specific site. The direct crushing of vegetation by trampling is one factor that favors the replacement of natural vegetation by non-native basal rosette- type plants such as plantain or hawkweed. Soil compaction caused by human treading retards the growth of trees, perhaps killing them. Forest duff can be pulverized, the soil denuded, the ground can become puddled and down-slope erosion can occur. The long-term effects of these impacts are visibly and seriously to alter the original vegetation patterns and associations ina manner that normal plant succession would not. Natural vegetation patterns contribute to the characterization of the unique or valued aspects of the particular site in the first place. Further- more, wildlife that requires special vegetational habitats will be affected. Campgrounds in some parks, for example, contain populations of Common Grackles, Brown-headed Cowbirds, and even House Sparrows. These are species not normally found in any numbers in the sur- rounding natural habitat. The non-consumptive user is guilty of these impacts. Point Pelee National Park in Ontario has been hammered by birdwatchers. The impact of camping has been so great there that it is no longer allowed. Restrictions are now being placed on the number of people entering some of our large semi-wilderness parks to keep down the collective damage (and to protect the visitors’ experience). North of Tofino, on the west coast of Vancouver Island, there are hot springs in Maquinna Provincial Park that are waded and bathed in by organized groups from hiking clubs. These rare hot springs are now ruined and valueless as an ecological reserve. Members of naturalist clubs are often the worst offenders in unique or highly sensitive habitats. These are areas we actively seek because of their high interest value. But we tramp around in bogs, marshes, alpine meadows, and gull colonies, content in our non-consumptive status. In- creasing numbers of natural food buffs are 1977 systematically harvesting edible wild nature. This problem is especially relevant in parks and similar reserves where the hunting or “har- vesting” of other wild things is forbidden, but where groups like Outward Bound teach live- off-the-land survival skills. The accumulation of garbage and litter in remote places is a very serious problem. Tons and tons of it are hauled out of our remote and accessible recreation areas every year (53 tons from the interior canoe routes of Algonquin Park, Ontario, in 1972 (Toronto Globe and Mail, 8 June 1974)). Imagine the garbage that piles up in the well known and easily reached areas. Garbage is not only unsightly, its presence can alter natural behavior patterns in some species of wildlife. We all know about bears and garbage, yet how can littering activity that leads to the destruction of “problem” bears be called non-consumptive? Not only is it necessary to remove portions of the original countryside initially to accommodate the non-consumers, but the impact that these users have on the remainder continues year after year to erode the landscape more. The massive numbers of such users, doing their collective “thing” on our natural landscapes, makes them (i.e., us!) the most consumptive and the most destructive of all groups of recreationists. So we are faced with an interesting irony: the “non- consumers” are shown to be the most serious consumers, simply by virtue of their numbers, by what they do, and where they do it. We must accept that the notion of non- consumptive use is a myth. There is simply no such thing as a non-consumptive user. After all, land use has implicit in it the idea of consump- tion. The idea of land use probably derives from the bizarre human misperception that all of non- human nature is merely a storehouse of re- sources. To say “non-consumptive use” 1s actually to speak a contradiction. The net result of all so-called non-consumptive recreational activity is the creation of a real scarcity of unimpaired environments. The increasing scarcity of unimpaired environments is proof of the gradual consumptive nature of our activities. Some may find it difficult to accept the position we have taken. After all, the idea of the non-consumptive user is firmly entrenched in WILKES: MYTH OF NON-CONSUMPTIVE USER 347 our vocabularies. He has been the celebrated mythical beast that we have often used to justify landscape preservation. Under the myth we have self-righteously pointed accusing fingers at other resource consumers. We can no longer hold that somehow we are better than they are simply because we think we are non-consumptive. With a new perspective we can approach old problems with a fresh and perhaps more fruitful outlook. Let us explore some possibilities. At least three implications come to mind if we are to reject the idea of non-consumptive uses. We must construct strict rules guiding our behavior when visiting natural landscapes. We must adopt a new attitude and approach to land- use planning as it applies to recreational land- scapes. These ideas are to some extent inter- related. Naturalist clubs must pay special attention to rules, or standards of conduct, in the outdoors. They often travel en masse to the most sensitive areas in their vicinity. We would recommend then that clubs make an effort to travel to special spots only very occasionally and when they do, they should travel in small groups. Choose places to go at a time of year when you'll do the least damage, and then stay on established pathways in small groups. Identify plants where they are, without picking bits off to check at home. We know a few “naturalists” who crash around looking for bird nests, and photographers who tear away the foliage for the proper camera angle. We do not say that nest records are unimportant, only that conscience often isn’t part of the equipment of the recorder. Each club should recognize for itself what measures are necessary to ensure the least consumption on the part of its members. Controls on behavior extend from the voluntary actions of clubs to the mandatory restrictions of government agencies. Nobody wants willingly to give up more liberties in a world in which they are rapidly eroding in all aspects, but it is time that naturalists begin a co- ordinated effort to get behavioral restrictions instituted, at least in our large semi-wilderness parks. Strict visitor controls appropriate in large parks include party size limits, the use of burnable containers only, and the use of stoves rather than fires where natural wood is at a 348 premium. The ideal situation would entail licensing all back-country users and regulating their numbers through a permit system. The licensing procedure has a double benefit. It allows agencies to know how many users there are, and it could mean a skills test prior to licensing. A skills test is very important because ignorant and unskilled people are using natural landscapes more and more, and they do the most damage. The prospect of licenses and permits for so- called “non-consumptive” users may make people recoil in horror. It conjures up the image of an enlarged bureaucracy to deal with it, as well as the spectre of more lost liberties. Back in the 1950s and 1960s, outdoor recreation of all sorts, but particularly in parks and equivalent reserves, was held to be a right, and available free to anyone. This idea has never been seriously challenged until now. The recreation we have been discussing is not a right any more; it is a privilege. We no longer live in the world of the 1960s. Solitude and wild nature are scarce. We would rather see the price for the privilege of using it paid in personal liberty than in the erosion of the unique character of the landscapes left to us. Strict controls will be made a widespread necessity anyway, when area by area, overuse becomes a crisis. If we have the vision to see that controls are necessary now, why don’t we have the courage and freedom to implement them? We have said something of the need for a new justification for landscape preservation. Total conservation is a four-part concern. Wisely managed use is just one part. Others are of equal weight and importance. They are preservation, restoration, and protection. Preservation figures importantly in overall conservation, and yet the rationale for the preservation of landscape is almost always that recreational benefits accrue to the using public. We have gone so far as to equate parks with “preserved” land, when nothing is further from the truth. If we reject the idea of the non-consumptive user, and yet recognize the importance of landscape preserva- tion, we can hardly endorse parks as the appropriate vehicle for preservation, because parks are justified and developed for their recreational potential. This does not mean we should reject the idea of THE CANADIAN FIELD-NATURALIST Vo. 91 parks, but rather encourage governments to become serious about their stated purpose of preserving unimpaired landscapes. Neither should we reject the idea of people in parks, because there are regulatory mechanisms avail- able to limit resource consumption by tourists and others. But we must dismiss the idea that landscapes, and the communities of life on them, can only be preserved in parks, and that the rationale of preservation is recreation. Ecolo- gical reserves and nature conservancies are a step in this direction, but so far they have succeeded in setting aside only limited areas. It simply will not do any longer to justify parks, reserves and sanctuaries in terms of the benefits to be derived for the “non-consuming” public. This form at once categorizes these landscapes as “resources” anyway, and makes their eventual exploitation for recreation an imperative. It is sad to think that any justification is necessary at all for landscape preservation. Butif it is, then we should hold that natural land- scapes should exist for their own sake: that their internal dynamics are fundamental engines of nature, fueled by the sun, and nurtured by the earth. We should hold that landscapes and their internal dynamics should be preserved solely because they are there, for their own sake, and because they have the right to exist (see Stone 1974 for a discussion of the notion of legal rights for non-human nature). We must not only reject the idea that nature exists solely for human benefit, we must also develop new planning tools that are not based on human utility. Government land agency planning proceeds along conventional lines, and clubs and federa- tions expend considerable effort criticizing the results. We can call it “systems,” or “master,” or “site” planning; but what these terms really mean is the planning of how to accommodate people on the landscape. In natural areas conventional planning merely orchestrates the systematic reversal of the principles of preservation. If we recognize the consumptive nature of all recrea- tional land uses, and are really concerned about landscape preservation, then we should reject conventional land-use planning in favor of non- use planning. A new theory of non-use planning can be generated from a thorough understanding of the nature of resource consumption by recreation- OT ists. It would involve the identification of physical carrying capacities on natural land- scapes through detailed inventories and sam- pling. It would center around strict controls on the numbers and behavior of participants in supposedly non-consumptive pursuits. It would place preservation as the top priority instead of use. Finally, it would emphasize that non-human nature exists for its own sake, and that the accommodation of people in it is not a matter of compromise but rather one of integration. In this article, a critical evaluation of the notion of non-consumptive use was made and found to be false. Some of the implications of rejecting the notion were also explored. It remains to decide what to do next. The accep- tance and implementation of the various issues raised here could mark a new era for conser- vation in Canada. We have a choice: either we take cognizance of the future of natural land- scapes and organize ourselves to meet it now, or we languish, comfortable in the hope that somebody will do something when the crisis comes. For us, the price of waiting is too high. Acknowledgments Helpful comments and suggestions were grate- fully received from Ron Seaborn, Betty McKin- non, Bristol Foster, Yorke Edwards, Mary Ann Wilkes, and Karen Dawe. Special credit is due to Mollie Byrne, Dave Denis, Al Hawryzki, and Neil Dawe for serving on the editorial com- WILKES: MYTH OF NON-CONSUMPTIVE USER 349 mittee. The inputs received from the member clubs of the Vancouver Island Region are also gratefully acknowledged. Literature Cited Bates, G. H.1935. The vegetation of footpaths, sidewalks, cart-tracks and gateways. Journal of Ecology 23: 470-487. Bouchard, A. 1973. Carrying capacity as a management tool for national parks. Park News 9(3): 39-52. Frissell, S.S. and D. Duncan. 1965. Campsite preference and deterioration in the Quetico-Superior Canoe Country. Journal of Forestry 63(4): 256-260. La Page, W. F. 1967. Some observations on campground trampling and ground cover response. United States Department of Agriculture Service, North East Forest Experiment Station, Research Paper N.E.-68. Lucas, R. C. 1964. Wilderness perception and use: The example of the Boundary Waters Canoe Area. Natural Resources Journal 3(3): 394-411. Meinecke, E. P. 1929. The effect of excessive tourist travel on the California Redwood Parks. California State Printing Office. 20 pp. Lime, D. W. and G. H. Stankey 1971. Carrying capacity: maintaining outdoor recreation quality. Jn Recreation Symposium Proceedings. Edited by E.V.H. Larsen. United States Department of Agriculture Forest Service. North East Forest Experiment Station, 1971. Stone, Christopher. 1974. Should trees have standing? Toward legal rights for natural objects. Kaufmann, Los Altos, California. Stone, G.P. and M.J. Taves. 1956. Research into the human element in wilderness use. Society of American Foresters Proceedings. pp. 26-32. Received 6 June 1977 Accepted 21 August 1977 Some New and Interesting Grass Records from Southern Ontario P. M. CATLING, A. A. REZNICEK, and J. L. RILEY Department of Botany, University of Toronto, Toronto, Ontario M5S IAI Catling, P. M., A. A. Reznicek, and J. L. Riley. 1977. Some new and interesting grass records from southern Ontario. Canadian Field-Naturalist 91(4): 350-359. Locations, and remarks on ecology, distribution, and taxonomy are given for certain grasses that represent either additions to the flora or significant range extensions. These grasses range from very rare native species of unusual habitats that should be protected, to aggressive weedy species that have been introduced from Eurasia. Included are Andropogon virginicus, Aristida basiramea, Aristida dichotoma, Aristida longispica, Aristida necopina, Aristida oligantha, Aristida purpurescens, Deschampsia cespitosa var. parviflora, Eragrostis spectabilis, Leptochloa fascicularis, Leptoloma cognatum, Muhlenbergia asperifolia, Panicum dichotomiflorum, Panicum sphaerocarpon, Panicum spretum, Panicum rigidulum, Poa bulbosa, Setaria faberi, Spartina patens, Tridens flavus, and Zoysia japonica. Key Words: grasses, Ontario, distribution, ecology, taxonomy, introduced, endangered, weeds. We here report as new to the province of Ontario 11 species. These are Andropogon virginicus, Aristida longispica, Aristida oli- gantha, Leptochloa fascicularis, Muhlenbergia asperifolia, Panicum spretum, Poa _ bulbosa, Setaria faberi, Spartina patens, Tridens flavus, and Zoysia japonica. Seven of these, Andropo- gon virginicus, Aristida longispica, Aristida oligantha, Leptochloa_ fascicularis, Setaria faberi, Tridens flavus, and Zoysia japonica are apparently new to Canada. Ten other rare species are also discussed. Of these 21 grasses, Andropogon virginicus, Aristida purpurescens, Panicum sphaerocarpon, Panicum spretum, and Panicum rigidulum are certainly among our rare native species and the status of some of these in Ontario may be precarious. The stations of these species should be considered for protection. Aristida longispica and Aristida necopina are probably also very rare, but they are so inconspicuous that further investigation of suitable habitats will be necessary to determine their status precisely. Deschampsia cespitosa var. parviflora, Leptochloa fascicularis, Muh- lenbergia asperifolia, Poa bulbosa, Setaria faberi, Spartina patens, and Zoysia japonica are clearly introduced species. Other species, notably Aristida basiramea, Aristida dichoto- ma, Aristida oligantha, Eragrostis spectabilis, Leptoloma cognatum, and Tridens flavus may well have been members of our original flora but some or all of our collections are from sites where introduction is a possibility. Our collection localities fall into two areas of Ontario, the Carolinian Zone (Soper 1962) and the eastern part of the Georgian Bay region in Muskoka District and Simcoe County. Records from the latter region include Aristida basi- ramea, Panicum sphaerocarpon, Panicum spretum, and Panicum rigidulum. Soper (1956) notes this area as a region where there is a concentration of southern species beyond the Carolinian Zone. The species are all listed here in alphabetical order and under each are specimen citations followed by a short discussion of the species’ occurrence with notes concerning ecology, general distribution, and taxonomy. All Ontario collections seen by the authors are listed except for Panicum dichotomiflorum and Panicum sphaerocarpon. Andropogon virginicus L. Norfolk County: Windham township, 4 mi (0.8 km) east of LaSallette, relict prairie. P. F. Maycock, O. B. Maryniak 5558, 20 May 1958 (sub. A. scoparius) (DAO, herb. P. F. Maycock). Welland County (Regional Municipality of Niagara): Bertie Township, open coal cinders on north side of railway yards south of Jarvis Street, about % mi (0.8 km) west of Fort Erie North. P. M. Catling, J. L. Riley, 23 October 1976 (CAN, DAO, TRT). Kent County: Zone township, “Thamesville Moor,” east side of Florence Road (County Road 26), about Y%—Y mi (0.4-0.8 km) north of Highway 21 and northwest of Thamesville Station. P. M. Catling, L. Cwynar, B. Freedman, 26 May 1977 (DAO, TRT). Broom-sedge, a native species with straight awns and smaller florets than A. scoparius, is found throughout much of the eastern and midwestern 350 SII United States (Hitchcock 1951). It is known from 11 counties in southern Michigan (Voss 1972) but has not yet been found in Ontario’s southwestern counties of Essex, Lambton, and Middlesex. The Niagara record cited above represents the first record for the Niagara Frontier region (Zenkert 1934; Zenkert and Zander 1975). The Norfolk County specimen is apparently the first Canadian collection of this species and was from a native prairie habitat. Aristida basiramea Vasey Norfolk County: about 1.6 mi (2.6 km) west of lighthouse, Long Point. J. E. Cruise 7803, 6 Sept- ember 1954(sub. A. intermedia) (DAO, TRT). Simcoe County: Tiny township, Lot 7, Concession XVIII, 44 mi (7.2 km) northwest of Penetang. A. A. Reznicek 4538, 20 August 1975 (DAO, TRT). The Norfolk County collection was reported as A. intermedia (Cruise 1969). Aristida basiramea has been reported repeatedly from Manitoba, the first report being that of Macoun (1888), but the reports apparently have not been confirmed (Scoggan 1957) and in Canada this species is still known conclusively only from Ontario. At the Simcoe County station A. basiramea is widely scattered and sometimes frequent in areas of dry bare sand in full sun, associating with Carex rugosperma, Panicum depauperatum, and Sporo- bolus cryptandrus. This station is 150 mi (245 km) farther north than any other Aristida record in Ontario and is the only occurrence of a species of this genus north of the Carolinian zone. In this regard it is interesting to note that A. basiramea is also the only species of Aristida to be found north of the “tension” zone in Michigan (Voss 1972). Aristida dichotoma Michaux Locality unknown: Three Pigeons, Canada. (NY, Photo DAO, Photo TRT). (This obscure, old record is probably not from Canada as the locality is untraceable). Welland County (Regional Municipality of Niagara): Port Colborne. J. C. Mc Rae, 1880(MTMG, Photo DAO). Regional Municipality of Niagara: Bertie township, railway yards, north side, south of Jarvis Street, about 4 mi(0.8 km) west of Fort Erie North. P. M. Catling, J. L. Riley, 23 October 1976 (CAN, DAO, TRT, TRTE). The Port Colborne collection represents the oldest record of an Aristida species in Ontario, and was noted by Macoun (1888). At the recently discovered Fort Erie Station this species occurs by the thousands over an extensive area on open and dry coal cinders, where it was associated with the somewhat less frequent Aristida oligantha. CATLING ET AL.: NEW GRASS RECORDS, SOUTHERN ONTARIO 351 Aristida longispica Poiret Kent County: Zone township, “Thamesville Moor,” east side of Florence Road (County Road 26), about Y—'4 mi (0.4-0.8 km) north of Highway 21 ai northwest of Thamesville Station. P. M! Catling, A. A. Reznicek, S. M. McKay, R. Brown, 18 Sept- ember 1976 (CAN, DAO, OAC, TRT, TRTE). This station represents the first Canadian record of this species. The plants were found in close asso- ciation with A. necopina in open sandy soil. Like other species of Aristida, this one begins to flower late in August and does not develop fruit until September or October. Even then it may be relatively inconspicuous. Aristida necopina Shinners (A. intermedia auct.) Norfolk County (mistakenly listed as Leeds Coun- ty) Longue Pointe, sur les sables découverts. F. Marie-Victorin, F. Rolland-Germain, E. Jacques, 8 aout 1932 (sub. A. gracilis) (DAO). Kent County: Zone township, “Thamesville Moor,” east side of Florence Road (County Road 26) about Y%,—'4 mi (0.4-0.8 km) north of Highway 21 and northwest of Thamesville Station. P. M. Catling, A. A. Reznicek, S. M. McKay, R. Brown, 18 Sept- ember 1976 (TRT). Zone township, about 2 mi (3.2 km) north of Thamesville Station, on the west side of County Road 26 (to Florence). P.M. Catling, A. A. Reznicek, S. M. McKay, R. Brown, 18 September 1976 (DAO, TRT, TRTE). The collection from Long Point in Norfolk County was reported as Aristida intermedia (Cruise 1969). The county given on the label of this collection is Leeds. This is clearly an error in label preparation since labels of other herbarium specimens indicate that Victorin et al. were on Point Pelee, Essex County, on 7 August 1932, and at St. Willams, Norfolk County, on 8 August 1932 (TRT). There are a number of Carolinian species in the 8 August Long Point collections which do not occur in Leeds County. Long Point in Norfolk County is only a few miles from St. Williams and well within the Carolinian zone. The recently discovered stations in Kent County are characterized by open sandy soil and an open prairie- like habitat with Danthonia spicata, Solidago nemo- ralis, and a haircap moss (Polytrichum sp.). At the Thamesville moor A. necopinaand A. longispica grew in close association. Aristida oligantha Michaux Welland County (Regional Municipality of Niagara): Bertie township, railway yards, north side, south of Jarvis Street, about '4 mi(0.8 km) west of Fort Erie North. P. M. Catling, J. L. Riley, 23 October 1976 (CAN, DAO, TRT, TRTE). Sa This is the first Canadian record. A few hundred plants were found in dry coal cinders in the open. Associates included other rare grass species such as Andropogon virginicus, Aristida dichotoma, Era- grostis spectabilis, and Leptoloma cognatum. Aristida purpurescens Poiret Essex County: Ojibway Prairie, Windsor, W. Botham, 8 September 1968 (herb. W. Botham). Dry prairie at Ojibway, south of Windsor, P. F. Maycock, 24 October 1969 (Photocopy DAO). South of Windsor, “LaSalle” prairie, west of Highway 18 behind Naminco Factory, P. W. Ball, P. F. Maycock 68271a, 9 September 1971 (TRTE). South of Windsor, Ojibway prairie, Matchette Road, south end, west side, P. W. Ball, P. F. Maycock 70471la, 10 September 1971 (TRTE). Windsor prairie, east of Windsor Raceway and Ojibway Park, P. M. Catling, S. M. McKay, 21 September 1975 (TRT). Dodge (1914) reported this species from dry ground on Squirrel Island in Lambton County, but it was not found there in Gaiser’s later survey (Gaiser and Moore 1966), nor has it been encountered there since. We have found it to be relatively frequent in the drier parts of the relict prairies south of Windsor but it was not recorded there by either Rogers (1966) or Thompson (1975). Aristida purpurescens becomes conspicuous only in late summer and autumn, and is usually not apparent by persistent remains, despite its perennial habit and relatively large size. Deschampsia cespitosa (L.) Beauv. var. parviflora (Thuill.) Coss. & Germ. Perth County: 4 mi (6.4 km) northwest of Stratford along Highway 8. W. G. Dore, H. A. Senn 47-430, 30 July 1947 (DAO, TRT). Carleton County: about 2 mi (3.2 km) above Billings Bridge, Ottawa, W. G. Dore 13600, 29 September 1951 (DAO). Simcoe County: Essex township, Lot 22, Concession VI, 444 mi (7.2 km) southeast of Angus. A. A. Reznicek 4533, 6 September 1975 (DAO, TRT). The widespread species Deschampsia cespitosa, represented by var. glauca (Hartm.) Lindm. f. 1s, throughout southern Ontario, a local and delicate species of calcareous shores and limestone plains. It is only rarely a weed of disturbed sites. Deschampsia cespitosa var. parviflora, introduced from Europe, is a locally aggressive colonizer of ruderal sites. It is a tall variety, frequently to | m, with large panicles, long flat leaves and small spikelets averaging 2.5-3.0 mm long with the first glume 1.9-2.7 mm and the second glume 2.3—2.6 mm (in the material cited above). This variety was reported as locally established in New England long ago (Fernald 1926). It is also THE CANADIAN FIELD-NATURALIST Vol. 91 known from the Maritime provinces (Roland and Smith 1963-64; DAO) and Quebee (Boivin 1967: DAO). The Carleton County station, probably result- ing from persistence in an old garden (Dore 1959), appears to have been destroyed (Dore, personal communication). An extensive stand was found in the ditch along Highway 8 near Stratford, and the Simcoe County station is similarly in moist open sandy ditches along a road. Deschampsia cespitosa var. parviflora is as yet very rare in Ontario. Eragrostis spectabilis (Pursh) Steudel Frontenac County: Sharbot Lake, 1.5 mi (2.4 km) southwest, about 50 clumps along railroad shoul- der. W. G. Dore 13871, 31 July 1952 (DAO). Oxford County: Woodstock, sandy ground, lawn of park on Ingersoll Avenue. F. H. Montgomery, W. Shumovich, 14 September 1953 (DAO, OAC, TRT). Kent County: open sandy ground, about 4 mi (6.4 km) north of Bothwell on north side of road west of railway crossing. P. M. Catling, A. A. Reznicek, S. M. McKay, R. Brown, 18 September 1976 (TRT). Welland County (Regional Municipality of Niagara): Bertie township, dry open cinders and gravel, north side of railway yards south of Jarvis Street, about % mi (0.8 km) west of Fort Erie North. P. M. Catling, J. L. Riley, 23 October 1976 (CAN, DAO, TRT). This is our only perennial Eragrostis species. It has a hard knotty base and large purplish inflorescence which is readily detached at maturity, acting as a tumbleweed. The inflorescence is larger than our annual members of the genus, 19-30 cm long and 15-26 cm wide in specimens from the locations cited above. Called “Tumble-grass,” “Purple Lovegrass” and, interestingly, “Petticoat-climber,” this grass was first reported from sandy beaches on Pelee Island (Core 1948). In addition to the collections cited above, it has also been reported for Elgin County (W. G. Stewart 1972, unpublished data) and Essex County (Thompson 1975). It is widespread in the lower peninsula of Michigan (Voss 1972), and throughout the eastern, midwestern, and part of the southwestern United States (Fernald 1950). Undoubtedly it will be found elsewhere in Ontario. The collections cited above represent the first records for Kent County and for the Niagara Frontier region (Zenkert 1934; Zenkert and Zander 1975). Elsewhere in Canada, EF. spectabilis is known only from southwestern Quebec (Boivin 1967). This grass may well be native in prairie-like habitats on the Bothwell sand plain in Kent County and in the, Windsor prairie (Thompson 1975). 1977 Leptochloa fascicularis (Lam.) Gray District Municipality of Niagara: Montrose railway yard, south side of Niagara Falls. P. M. Catling, K. L. McIntosh, 27 September 1976 (CAN, DAO, TRT, TRTE). Salt Meadow Grass is found in brackish marshes along the Atlantic coast and on alkaline flats in western North America (Hitchcock 1951), but is rare inland in the east and has not previously been reported for Canada. It occurs inland in New York State mostly in the vicinity of salt springs and salt factories (Wiegand and Eames 1925) but also in railway yards (Zenkert and Zander 1975). It has also been found in Michigan (Stephenson 1967). The occurrence of this species in railway yards is usually associated with the stock- piling or transport of salt, and with the use of salt to melt ice and snow on the switches and elsewhere in the yards. The plants cited above have the second glume 3.54.3 mm long and awns 0.9—1.2 mm long. They are referable to var. acuminata(Nash) Gleason, which has western affinities (Gleason and Cronquist 1963). This grass has been frequently called Diplachne acuminata (Fernald 1950). Leptoloma cognatum (Schultes) Chase Peel County: prairie remnant beside railway, Lorne Park. P. W. Ball, 22 September 1971 (TRTE). Welland County (Regional Municipality of Niagara): Bertie township, dry open cinder and gravel, north side of railway yards, south of Jarvis Street, about Y% mi (0.8 km) west of Fort Erie North. P. . Catling, J. L. Riley, 23 October 1976 (CAN, DAO, TRT, TRTE). Widely distributed in southern Michigan (Voss 1972) and scattered through New York State (Zenkert and Zander 1975), it appears that the records cited above represent the second and third records for the province. It was first reported in Ontario from Rockhouse Point in Haldimand County (Zenkert and Zander 1975). Fall Witchgrass occurs throughout most of the eastern and midwestern United States. It is apparently native to the west and south, but may be only adventive in the northeast, where it has a more scattered distribution (Hitchcock 1951; Gleason and Cronquist 1963). From a distance this grass resembles Panicum capillare or Muhlenbergia asperifolia. The mem- branous ligule, nearly obsolete first glume, and long second glume, and the narrowly elliptic shape of the spikelets are helpful in recognition. Muhlenbergia asperifolia (Nees and Meyen) Parodi Essex County: east of Windsor Salt factory, Windsor. CATLING ET AL.: NEW GRASS RECORDS, SOUTHERN ONTARIO 353) P. M. Catling, S. M. McKay, 18 August 1975 (CAN, DAO, TRT). District Municipality of Niagara: south side of Niagara Falls, Montrose railway yard. P. M. Catling, K. L. McIntosh, 27 September 1976 (CAN, DAO, TRT). In both of the above localities the plants were found in several large patches on open and highly alkaline soil. These are the first Ontario records and may be the first records for northeastern North America. Muh- lenbergia asperifolia is found in alkaline prairies, sloughs and ditches in western North America. Hitchcock (1951) lists New York as part of its range, but does not map this species east of Indiana. It has not been reported from Michigan (Voss 1972). Elsewhere in Canada this species is found from southern Manitoba to British Columbia (Scoggan 1957; Boivin 1967). Panicum dichotomiflorum Michaux Peel County: Toronto township (city of Mississauga), Y% mi (0.8 km) south of junction of Dundas Street (Highway 5) and Mississauga Road, on south bank of Credit River. A. A. Reznicek, D. R. Gregory, 27 October 1975 (TRTE). Welland County (Regional Municipality of Niagara): Grantham township, open alkaline median of the Queen Elizabeth Highway at St. Catharines. P. M. Catling, K.L. McIntosh, 10 September 1976 (TRT). (Regional Municipality of Niagara): Niagara Falls, open alkaline soil, Montrose railway yard on south side of town. P. M. Catling, K. L. McIntosh, 27 September 1976 (TRT). (Regional Municipality of Niagara): Bertie township, railway yards north side, south of Jarvis Street, about '4 mi(0.8 km) west of Fort Erie North. P. M. Catling, J. L. Riley, 23 October 1976 (CAN, DAO, TRT). York County: East Toronto, open soil along a zinc fence, Lakeshore and Leslie intersection. P. M. Catling, B. Freedman, 24 October 1976 (CAN, DAO, TRT). Although a native of eastern North America, this late-flowering annual has weedy tendencies. It appears to have spread into rural and cultivated lands in extreme southwestern Ontario about 1920. Until recently it was considered a relatively un- common plant known only from the southwest and from the far eastern parts of Ontario in the Ottawa and St. Lawrence lowlands where it was first noticed in 1949. Only recently was it reported from the Canadian portion of the Niagara Frontier region (Yaki 1970; Zenkert and Zander 1975). Our recent field work has disclosed that this species is now quite 354 frequent and, in fact, a serious weed of cornfields in the extreme southwest. Its weedy nature has been documented by Alex and Switzer (1976). Panicum dichotomiflorum, or at least this ecotype (which may be of recent origin), is a tough plant tolerant of extremely dry, exposed habitats with levels of certain metals in the soil that would be toxic to many plants. Not surprisingly it appears to be spreading. Most of the specimens cited above may be clearly segregated as var. geniculatum (Wood) Fern. Panicum rigidulum Nees Hastings County: about | mi(1.6 km) south of Stoco. W. G. Dore 13490, 15 August 1951. (sub. P. agros- toides) (DAO). On shore of Moira River at the bridge, Stoco. W.G. Dore 13493, 15 August 1951 (sub. P. agrostoides) (DAO). Wet rocky flood-shore of bay of Stoco Lake, Tweed. W. G. Dore 13496, 15 August 1951 (sub. P. agrostoides) (DAO). Muskoka District: Wood township, Concession VI, Lot 37, Hurling Point, | mi (1.6 km) west of Bala. R. E. Whiting, L. Jewitt 1437, 23 August 1972 (sub. Lei THE CANADIAN FIELD-NATURALIST Vol. 91 P. agrostoides) (herb. R. E. Whiting, TRT). Morrison township, VIII-31, northeastern shore of Sparrow Lake, 4.3 km west southwest of Kilworthy Station. A. A. Reznicek, R. Bobbette 2391, 7 Sept- ember 1975 (herb. R. E. Whiting, TRT). This very rare species is apparently native on rocky shores near the southern edge of the Canadian Shield (Figure 1). The two recent Muskoka collections are both from crevices in Precambrian rock directly on the shores of lakes. They were not widely separated and the species is likely scattered on other, similar shores in this region. The plant is reported by Stroud (1941) from Wellington County, Ontario, apparently based on a specimen alleged to be from Guelph: “meadows and pastures, Guelph, Ontario, L. McDougall, July 1937” (DAO). It is very likely that the specimen has been mislabelled as per annotations on the sheet by W. G. Dore and B. Boivin. Panicum rigidulum is unknown elsewhere is eastern Canada, but was reported long ago from British Columbia (Macoun 1888). It is known from five counties in southern Michigan (Voss 1972) and is reported from North Bass Island in western Lake Erie (Core 1948). This would seem to make its presence in extreme southwestern Ontario a possibility. PIS) ake FIGURE |. Wet rocky flood-shore habitat of Panicum rigidulum. Photographed at Stoco Lake, near Tweed, Hastings County, 15 August 1951 (DAO), by W. G. Dore. SIT This species is known in many floras (Fernald 1950; Gleason 1952) as P. agrostoides, a fitting but illegitimate name (Voss 1966). In view of the fact that we also discuss the occurrence of Leptoloma cognatum in Ontario, it should be noted that one of the key characters often used to separate Leptoloma from Panicum is the membranaceous ligule of Leptoloma as contrasted with the ring of hairs or obsolete ligule of Panicum (Shinners 1944; Fassett 1951). Panicum rigidulum, however, has a distinctive, membranaceous, erose ligule quite different from other Ontario Panicum species. Panicum sphaerocarpon Ell. Simcoe County: North Orillia township, Concession X, Lot 24, Matchedash Lake, 10 mi (16 km) northeast of Coldwater. A. A. Reznicek 4543, 27 August 1976 (DAO, TRT). Kent County: Zone township, “Thamesville Moor,” east side of Florence Road, County Road 26, about Y%—Y mi (0.4-0.8 km) north of Highway 21, north- west of Thamesville Station. P. M. Catling, A. A. Reznicek, S. M. McKay, R. Brown, 18 September 1976 (DAO, TRT). This infrequent southern species of moist-to-dry sandy sites was known froma number of collections in Essex, Kent, Lambton, and Middlesex Counties in southwestern Ontario. It was also reported for Elgin County by W. G. Stewart (1972, unpublished data). Our second collection adds a site in northern Kent County near the Middlesex County border. In addition, the collection from Simcoe County rep- resents a range extension of about 170 mi (275 km) from the nearest Ontario site for the species. It occurs here on the sandy shores of Matchedash Lake with a host of other interesting species as described in the article under Panicum spretum. Panicum sphaerocarpon does not occur north of the “tension” zone in Michigan (Voss 1972). Thus, the Simcoe County disjunction is probably real and not an artifact due to lack of collecting of Panicum. Panicum sphaerocarpon has nevertheless been over- looked in parts of southern Ontario. We have found it at several additional locations in Kent County, and we were surprised to find that its occurrence had not been documented at any of these sites before. Panicum sphaerocarpon is quite distinctive in its interesting juxtaposition of very small spikelets with wide leaves, up to 19 mm in the Simcoe County material. The leaves of the conspicuous fall rosettes are especially broad with many strong nerves. Panicum spretum Schultes Simcoe County: North Orillia township, Lot 24, Concession IX, Matchedash Lake,9!4 mi(15.3 km) CATLING ET AL.: NEW GRASS RECORDS, SOUTHERN ONTARIO 355 northeast of Coldwater. A. A. Reznicek, 30 June 1975 (DAO). Matchedash Lake, P. W. Ball 26575, 30 June 1975 (TRTE). Matchedash township, Lot 1, Concession VIII, Matchedash Lake, 8 mi (12.9 km) northeast of Coldwater. A. A. Reznicek 4544, 13 August 1976 (TRT). Muskoka District: Wood township, Lot 16, Con- cession XIV, Nine-mile Lake. P. F. Maycock, 23 October 1976 (herb. P. F. Maycock, Erindale College). This primarily coastal-plain species was first found in Ontario on drying sandy lakeshores in Simcoe County in 1975 and subsequently in nearby Muskoka District in 1976. It is one of the coastal plain species discussed by Voss (1972) who noted that it is very rare in Michigan. The only other Canadian sites of this species are in southern Nova Scotia (Dore and Roland 1942; Roland and Smith 1969). When robust, the species may be up to | m tall and is unmistakable on the sandy lakeshore. Panicum spretum may be distinguished from related species of Panicum in the region by its size, glabrous foliage and sheaths, long ligule and narrow panicle. In its sandy lakeshore habitat, Panicum spretum occurs with other rare plants, some of which are also of eastern and coastal affinity. These include Rhexia virginica, Linum striatum, Muhlenbergia uniflora, Fimbristylis autumnalis, Panicum sphaerocarpon, Rhynchospora capitellata, Polygonum careyi, and Xyris difformis. Poa bulbosa L. Peel County: Mississauga township, several scattered clumps along 10 yards of steep, recently disturbed sandy bank, north side of Highway 5, east of Mississauga Road, community of Erindale. A. A. Reznicek 3778, 27 May 1972 (DAO, TRT, TRTE). Mississauga township, roadside, Erindale, P. W. Ball 2132, 14 May 1973 (TRTE). Mississauga township, dry gravelly roadside bank, southwest corner of interchange of Queen Elizabeth Highway and Mississauga Road, Mississauga. A. A. Reznicek 4545, 29 May 1975 (TRT, TRTE). York County: West Toronto, edge of disturbed path to woods, off Royal York Road, north of bridge, southwest edge of Lambton Woods. E. Hamilton, 3 June 1976 (TRT). Bulbous Bluegrass, introduced from Europe, is a very distinctive species with the florets converted to bulblets with dark purple bases. It is widespread in western North America (Hitchcock and Cronquist 1973) and in Canada, frequent in the Victoria region of British Columbia (Szczawinski and Harrison 1973). In eastern North America, it is very local and 356 scattered, having been reported from New York, Virginia, and North Carolina (Hitchcock 1951). These are the first collections of this species growing without cultivation in Ontario. The records are all from the Toronto region, and may indicate a developing local infestation. The collections cited above may be referred to P. bulbosa var. vivipara Koeler. A specimen at OAC labelled “waste places; Kemptville, Ontario; 1939” is believed to be mis- labelled (W. G. Dore, personal communication). Setaria faberi Herrm. District Municipality of Niagara: Montrose railway yard, south side of Niagara Falls. P. M. Catling, K. L. McIntosh, 27 September 1976 (CAN, OAC, TRT). Welland County (Regional Municipality of Niagara): Bertie township, north side of railway yards south of Jarvis Street, about 14 mi (0.8 km) west of Fort Erie North. P. M. Catling, J. L. Riley, 23 October 1976 (CAN, DAO, TRT). York County: Metropolitan Toronto, alonga railway, Don Valley, north of Bloor viaduct, opposite brickyard. P. M. Catling, J. Kaiser, S. M. Mckay, 28 September 1976 (CAN, DAO, TRT). Metropolitan Toronto, along a railway track, south of Wicksteed Avenue, East York. P. M. Catling, J. Kaiser, K.-L. McIntosh, S. M. McKay, | October 1976 (CAN, DAO, OAC, TRT). East Toronto, along C.N. railway tracks in the railway yard between Victoria Park Avenue and Main Street, near old Danforth Station. P. M. Catling, S. M. McKay, K. L. McIntosh, 3 October 1976 (CAN, TRT). : These are the first records for Ontario for this introduced Asian species. It is spreading aggressively in the northeastern and midwestern United States (Reed 1970). In Ontario, it occurs now as one of a characteristic group of plants, such as Plantago psyllium and Chaenorrhinum minus that are found mainly in gravel and/or coal cinders and clay in open locations along railways, and especially in railway yards (Figure 2). This species resembles S. viridis but differs in having a strongly nodding panicle (up to 14 cm long) bent below the middle, spikelets mostly over 2.5 mm long, the fertile lemma more or less tapering to a distinctly exposed tip, and leaf blades more or less hairy above (Voss 1972). In addition, it is tetraploid while S. viridis is diploid (Pohl 1962). Pohl (1962) has also pointed out that the blades are not always hairy, but in all of our material cited above the upper leaf blades are distinctly hairy. In depauperate plants the panicles may not be long enough to curve over, and are often quite erect. Also, depauperate specimens may have only one flower per panicle branch, giving THE CANADIAN FIELD-NATURALIST Vol. 91 FiGuRE 2. A patch of Setaria faberi with characteristic nodding, spike-like panicles. These plants were growing in cinders and gravel along a railway track at Fort Erie North, District Municipality of Niagara. Photographed 23 October 1976 by P. M. Catling. the impression of as many as 10 spines per floret only because the other florets have not developed. The latter phenomena also occurs towards the base of longer spikes, so that the use of bristle number is an unreliable identification feature. Even in depauperate material of S. faberi, the short second glume covers only three quarters to five sixths of the fertile lemma, instead of 9/10 asin S. viridis, and the prominent cross- ridges of the fertile lemma contrast with the minutely papillose lemmas of S. viridis (often spotted with dark brown). Depauperate plants of S. faberi(to 20 cm tall) have spikelets 3 mm long compared to similarly sized plants of S. viridis with spikelets only 2 mm. Spartina patens (Ait.) Muhl. Essex County: City of Windsor, immediately south of Windsor Salt factory (Sandwich Avenue and Propect Avenue). P. M. Catling, S. M. McKay, 17 May 1975 (CAN, DAO, TRT). Windsor Salt factory, Windsor. P. M. Catling, S. M. McKay, 21 September 1975 (CAN, DAO, TRT). 1977 These represent the first records of Salt-meadow Cordgrass for Ontario. The plants were found with other halophytes such as Solidago sempervirens and Juncus gerardii in open alkaline soil. Spartina patens is a characteristic species of salt marshes along the Atlantic coast with isolated inland stations previously known in New York and Michigan (Hitchcock 1951; Mobberley 1956; Voss 1972) where, like Leptochloa fascicularis, it occurs about sodium-salt springs, salt factories, and railway yards. Tridens flavus (L.) Hitche. Regional Municipality of Niagara: Montrose railway yard, south side of Niagara Falls. P. M. Catling, K. L. McIntosh, 27 September 1976 (CAN, DAO, TRT). Purpletop is widespread in the eastern and mid- western United States (Hitchcock 1951) and has long been expected in Ontario, this being the first record. It may have been formerly native in the Niagara peninsula and in extreme southwestern Ontario. It has not been reported elsewhere in Canada. A tall (1-1.5 m) attractive species, Purpletop may be readily overlooked owing to its superficial simila- rity to Panicum virgatum L. It differs in having the deep purplish spikelets 6- to 8-flowered. This grass is sometimes placed in the genus Triodia (Fernald 1950; Gleason 1952). Zoysia japonica Steudel Essex County: Mersea township, Point Pelee, W. Botham, 11 June 1972 (herb. W. Botham). Point Pelee National Park, just north of the Nature Interpretation Centre. A. A. Reznicek, P. F. May- cock, D. R. Gregory, 13 July 1976 (TRT, TRTE). This Asiatic species, and cultivars of it, have been recorded from lawns several times in the past from Ottawa, Windsor, and Kitchener (various collections, DAO and OAC). In these cases, it would appear to have been part of the sod or seed used to establish the lawns. In many instances Zoysia is an undesirable lawn grass. In Ontario at least it remains brown much later into the spring than other species and turns brown much earlier in the autumn. The collection cited from Point Pelee was not associated with a lawn. It grew in openings in a Juniperus virginiana glade on dry, sandy soil with Poa compressa, Sporobolus cryptandrus, Aster azureus, Euphorbia corollata, Rhus typhina, Opuntia com- pressa, and Cornus racemosa. The Zoysia in this colony had flowered well, in comparison with the vegetative condition of almost all other herbarium collections.. Although there were formerly cottages in the vicinity, this collection was not associated with their grounds and may well represent a natural establishment rather than a planting. CATLING ET AL.: NEW GRASS RECORDS, SOUTHERN ONTARIO 357 Acknowledgments We thank W.G. Dore and J. McNeill for considerable help with records of rare species and for allowing us to consult their manuscript on Ontario grasses (1977). B. Boivin helped with herbarium records. We also here express our gratitude to P. W. Ball, W. Botham, P. F. Maycock, and R. E. Whiting for allowing us to cite some of their recent collections. R. Brown kindly showed us some very interesting locali- ties in his local area of Kent and Lambton Counties, and this led to a number of dis- coveries. Finally, S. McKay and K. McIntosh both shared in the discoveries of several species, and for their assistance we are very grateful. Literature Cited Alex, J.F. and C.M. Switzer. 1976. Ontario weeds. Ontario Ministry of Agriculture and Food, Publication 505. 208 pp. Boivin, B. 1967. Enumération des plantes du Canada. Naturaliste Canadien 94; 471-528. Core, E. L. 1948. The flora of the Erie islands. Ohio State University, Franz Theodore Stone Laboratory Con- tribution 9. 106 pp. Cruise, J. E. 1969. A floristic study of Norfolk County, Ontario. Transactions of the Royal Canadian Institute 35: 1-116. Dodge, C. K. 1914. The flowering plants, ferns and fern allies growing without cultivation in Lambton County, Ontario. Michigan Academy of Science Annual Report 16: 132-200. Dore, W. G. 1959. Grasses of the Ottawa district. Canada Department of Agriculture Publication 1049. 101 pp. Dore, W.G. and J. McNeill. 1977. Grasses of Ontario. Agriculture Canada, Ottawa (In press.) Dore, W. G. and A. E. Roland. 1942. The grasses of Nova Scotia. Proceedings of the Nova Scotian Institute of Science 20: 177-288. Fassett, N.C. 1951. Grasses of Wisconsin. University of Wisconsin Press, Madison, Wisconsin. 173 pp. Fernald, M.L. 1926. Two summers botanizing in New- foundland. Rhodora 28: 145-156. Fernald, M. L. 1950. Gray’s manual of botany. 8th edition. American Book Company, New York. Ixiv + 1632 pp. Gaiser, L.O. and R.J. Moore. 1966. A survey of the vascular plants of Lambton County, Ontario. Plant Research Institute, Canada Department of Agriculture, Ottawa. 122 pp. Gleason, H. A. 1952. The new Britton and Brown illus- trated flora of the northeastern United States and adjacent Canada. New York Botanical Garden, New York. 3 volumes. Gleason, H. A. and A. Cronquist. 1963. Manual of vas- cular plants of northeastern United States and adjacent Canada. Van Nostrand, Princeton. li + 810 pp. Hitchcock, A. S. 1951. Manual of the grasses of the United States, 2nd edition. Revised by A. Chase. United States 358 THE CANADIAN FIELD-NATURALIST Department of Agriculture, Miscellaneous Publication 200. 1051 pp. Hitchcock, C.L. and A. Cronquist. 1973. Flora of the Pacific Northwest, an illustrated manual. University of Washington Press, Seattle. 730 pp. Macoun, J. 1888. Catalogue of Canadian plants. Part IV. Endogens. Dawson Bros., Montreal. 248 pp. Mobberley, D. G. 1956. Taxonomy and distribution of the genus Spartina. lowa State College Journal of Science 30: 471-574. Pohl, R. W. 1962. Notes on Seraria viridis and S. faberi (Gramineae). Brittonia 14: 210-213. Reed, C. F. 1970. Selected weeds of the United States. Agriculture Research Service, United States Department of Agriculture, Agriculture Handbook 366. 463 pp. Rogers, C. M. 1966. A wet prairie community at Windsor, Ontario. Canadian Field-Naturalist 80(4): 195-199. Roland, A. E.and E. C. Smith. 1963-64. The flora of Nova Scotia. Part I, Pteridophytes, Gymnosperms and Mono- cotyledons. Proceedings of the Nova Scotian Institute of Science 26(2): 3-238. Scoggan, H. J. 1957. Flora of Manitoba. National Museum of Canada Bulletin 140. 619 pp. Shinners, L.H. 1944. Notes on Wisconsin grasses. !V, Leptoloma and Panicum. American Midland Naturalist 32: 164-180. Soper, J. H. 1956. Some families of restricted range in the Carolinian flora of Canada. Transactions of the Royal Canadian Institute 31: 69-90. Soper, J. H. 1962. Some genera of restricted range in the Carolinian flora of Canada. Transactions of the Royal Canadian Institute 34: 2-56. Addendum Andropogon virginicus L. Kent County: Camden township, Lot 2, Concession II, about 2 mi (3.2 km) west of Dresden, P. M. Catling and R. Brown, 16 August 1977 (DAO, TRT). Howard township, on north side of Highway 401 at Interchange 13, about 6mi (9.6 km) south -of Thamesville in Concession VI, P. M. Catling, K. L. McIntosh and S. M. McKay, 27 August 1977 (DAO, TRT). Zone Township, Lot 6, Concession VIII, northwest side of Highway 2, about 3 mi(4.8 km) southwest of Bothwell, P. M. Catling and R. Brown, 27 August 1977 (DAO, TRT). Since it was first noticed in Ontario in 1976, this grass has been found at several localities on the Bothwell sand plain in Kent and Middlesex Counties. A selection of new localities is cited above. Vol. 91 Stephenson, S. N. 1967. Two additional grass species for southwestern Michigan. Michigan Botanist 6: 24-26. Stroud, J.J. 1941. A study of the flora of Wellington County, Ontario. Canadian Field-Naturalist 55: 56-62. Szczawinski, A. F. and A. S. Harrison. 1973. Flora of the Saanich Peninsula. British Columbia Provincial Museum, Occasional Papers 16: 1-114. Thompson, P.W. 1975. The floristic composition of prairie stands in southern Michigan. Jn Prairie: A. multiple view. Edited by M. K. Wali. University of North Dakota Press, Grand Forks, North Dakota. pp. 317-331. Voss, E.G. 1966. Nomenclatural notes on Monocots. Rhodora 68: 435-463. Voss, E. G. 1972. Michigan flora. Part I, Gymnosperms and Monocots. Cranbrook Institute of Science and University of Michigan Herbarium. Bloomfield Hills, Michigan. 488 pp. Wiegand, K.M. and A.J. Eames. 1925. Flora of the Cayuga Lake basin. Cornell University, Ithaca, New York. 491 pp. Yaki, G. J. 1970. Plants of the Niagara Peninsula. Niagara Falls Nature Club, Special Publication 2. 44 pp. Zenkert, C. A. 1934. The flora of the Niagara Frontier region. Bulletin of the Buffalo Society of Natural Sciences 16: 1-328. Zenkert, C. A. and R.H. Zander. 1975. Flora of the Niagara Frontier region, supplement. Bulletin of the Buffalo Society of Natural Sciences 16(sic): 1-62. Received 17 March 1977 Accepted 23 July 1977 Crypsis (Heleochloa) schoenoides (L.) Lam. Essex County: City of Windsor, south of the salt factory near Prospect Ave. and Sandwich Ave., P. M. Catling and K. L. McIntosh, 20 Sept. 1977 (DAO, TRT). A native of the Mediterranean region, this grass is known from various parts of the northeastern U.S. (Hitchcock 1951; Gleason and Cronquist 1963). It was first discovered in Ontario by J.W. Wilson of Windsor during the late summer of 1977. About 50 plants were found in moist organic soil (pH 7.5) of a ditch south of the Windsor salt factory. The plants grew in associa- tion with Cyperus odoratus, Pluchea purpurascens, and Spartina patens. Leptoloma cognatum (Schultes) Chase Kent County: Zone township, Lot 5, Concession III, along the Canadian Pacific Railway west of 1977 Highway 23, P.M. Catling and R. Brown, 16 August 1977 (DAO, TRT). At least 50 plants of Fall Witchgrass were found here in dry open sandy ground (pH 5.6) associating with Ambrosia psilostachya, Solidago nemoralis, Danthonia spicata, and Eragrostis spectabilis. Vhis represents the first record for Kent County and the fourth record for Ontario. Setaria faberi Herrm. Kent County: Dover township, north side of the mouth of the Thames River near Bradley’s Farms, P. M. Catling, 16 August 1977 (TRT). Camden township, Lot 2, Concession II, about 2 mi (3.2 km) west of Dresden, P. M. Catling and R. Brown, 27 August 1977 (DAO, TRT). CATLING ET AL.: NEW GRASS RECORDS, SOUTHERN ONTARIO 359 Chatham township, Lot 30, Concession I, about 4 mi (6.4 km) west of Dresden at Turner’s Pond, P. M. Catling and R. Brown, 27 August 1977 (DAO, TRT). Previously we stated that in Ontario this plant belongs to a group of species encountered mainly along railways. This is certainly true for parts of the province, but during our recent explorations of Kent County it was frequently found along the edges of fields of tomatoes, corn, and soybeans (specimens cited above). Here it associates with other weedy species such as Setaria viridis, Abutilon theophrasti, Panicum capillare, Solanum nigrum, and Chenopo- dium album. Received and accepted 12 October 1977 Terrestrial Oligochaeta of Some New Brunswick Caves with Remarks on Their Ecology DONALD F. MCALPINE! and JOHN W. REYNOLDS? 'Department of Natural Science, The New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K IES 2Department of Forest Resources, University of New Brunswick, Fredericton, New Brunswick E3B 5A3 McAlpine, Donald F. and John W. Reynolds. 1977. Terrestrial Oligochaeta of some New Brunswick caves with remarks on their ecology. Canadian Field-Naturalist 91(4): 360-366. Cave faunas are poorly known in eastern Canada. An examination of six caves and one abandoned mine in three New Brunswick counties revealed that four of the caves and the mine harbored earthworms (Annelida, Clitellata, Oligochaeta, Lumbricidae). Most frequently encountered were Dendrodrilus rubidus (Savigny) and Aporrectodea tuberculata (Eisen). Dendrodrilus rubidus seems to be the common cave earthworm in Canada as wellas inthe United States. Both of these species seem to be reproducing in the mine examined. There is no indication whether the other two species collected, Eisenia rosea (Savigny) and Lumbricus terrestris L., are established as troglophiles. The initial introduction of earthworms to caves or mines may have been accidental, e.g., through the actions of weather or man, but active colonization may play as important a role. The structure of the entrance to the cave or mine probably controls the primary method of earthworm introduction. Les faunas caves sont trés peu connues a |’Est du Canada. Une recherche dans six caves et dans une mine abandonnée dans trois comtés du Nouveau-Brunswick a révélé que quatre des caves et la mine sont hébergées de vers de terre (Annelida, Clitellata, Oligochaeta, Lumbricidae). Les plus frequemment trouvés furent Dendrodrilus rubidus (Savigny) et Aporrectodea tuberculata (Eisen). Dendrodrilus rubidus semble étre le plus commun des vers de terre des caves au Canada et méme aux Etats-Unis. Ces deux espéces semblent se propager dans la mine examinée. I] n’y a aucune indication que les deux espéces collectionnées, Eisenia rosea (Savigny) et Lumbricus terrestris L., sont établies comme troglophiles. L’introduction initiale des vers de terre dans les caves et dans les mines peut étre accidentel soit par des actes de la température ou de Phomme. La colonisation active peut aussi jouer un réle important. La structure de l’entrée de la cave et de la mine contréle probablement la méthode primaire de l’introduction des vers de terre. Key Words: caves, New Brunswick, ecology, distribution, earthworms, Aporrectodea tuberculata, Dendrodrilus rubidus, Lumbricus terrestris. Previous faunal investigations of Canadian caves have only briefly made mention of earth- worms. Halliday and Dunnet (1966) note Den- drodrilus rubidus (Savigny) was collected in Skull Creek Cave on Vancouver Island. (In a reprint of their paper a typographical error in the synonym used for the specific name of the species has rubida recorded as tubida. In noting this record the error was repeated by Peck and Fenton (1973) in a summary paper on the fauna of Canadian caves.) Calder and Bleakney (1965) found earthworms uncommon in their study of the microarthropod ecology of Frenchman’s Cave, a porcupine-inhabitated gypsum cave in Nova Scotia. The specimens they collected were not identified (Calder and Bleakney 1967). Recently, Reynolds (1977) has recorded D. rubidus, one of the 19 Ontario species, from mines in that province. It appears that the occurrence of Aporrectodea tuberculata (Eisen) in New Brunswick caves, recorded here, is the first published record of this species in such a habitat in North America. In many ways this may be owing to the past taxonomic problems in the trapezoides complex which have only recently been unravelled by Gates (1972). Gates (1959) reported A. twberculata may have been recorded from caves in Germany as A/lolobo- phora caliginosa (Savigny). Although from the subsurface habitat earth- worms normally occupy they would appear pre- adapted to the cave environment, they have not been one of the more successful troglobite3 groups. Vandel (1966) noted that troglobic terricoles have been described from Japan and Assam and recently Zicsi (1974) described a cavernicolous lumbricid from a _ subaquatic habitat in Hungary. No troglobic terricoles have been recorded from Canadian or American caves, though a number of limnicoles have been recently described from caves in the United States (Cook 1975). All earthworms in regions glaciated during the 3Animals that normally complete their life cycle only in caves; often showing specialized adaptations for cave life. 360 1977 Quaternary were exterminated and all mega- driles (= terrestrial oligochaetes) now present in these regions have been recently introduced by man (Gates 1970; Reynolds 1975, 1976a). There- fore, troglobic terricoles could not have evolved in the short time period since first European colonization of New Brunswick. Caves in this province appear too shallow to have served as refugia during the glacial period. As trogloxenic‘ and troglophilic,> earthworms have been more successful. Gates (1959) re- corded 12 species from caves in the United States but gives no indication whether actual popula; tions were involved, stating only that those. earthworms collected were surface forms. Most of the forms that Gates (1959) examined were parthenogenetic. a character considered favor- able to the colonization of a new area (Reynolds 1974). Gates (1959) noted that a single cocoon of some obligatory biparental species may give rise to a number of individuals. He felt that this, and the fact that copulation may be completed prior to the introduction into the new area, might indicate something other than that the repro- ductive mode may be of primary importance in the establishment of troglophilic earthworm populations. Nicholas (1960) has noted the difficulty in determining the extent of cave adaption ex- hibited by earthworms. Methods Investigations were carried out in six caves and one mine in Albert, Kings, and Saint John Counties of New Brunswick. All specimens were collected by hand-sorting through soil, detritus, gravel, and dung samples. Rocks and pieces of wood were overturned and sections of tree trunk and mine support beams were often split and examined. Details on habitat and zonation (dark or threshold zone) were recorded. Results and Discussion Four of the six caves and the mine were occupied by earthworms. From the 27 specimens collected, we identified 4 of the 14 species present in this province (Reynolds 1976b; Reynolds and 4Animals that frequently enter caves but do not complete their life cycle in caves. 5Animals that may complete their life cycle in caves but normally do so in suitable habitats outside of caves. MCALPINE AND REYNOLDS: OLIGOCHAETA ECOLOGY, NEW BRUNSWICK CAVES 361 Christie 1977). A number of the worms collected were too young to be identified to species. These all belonged to the genus Aporrectodea and are probably tuberculata since this was the only species of this genus obtained from the New Brunswick caves. Aporrectodea tuberculata is also the dominant earthworm species in Canada (Reynolds 1977). Table | records the data on the earthworm species collected at each location sampled. Earthworms were not abundant in any of the caves or mine (x = 5.4). Although no attempt was made to assess the total number of worms that each cave harbored, seven specimens was the maximum number of worms obtained by hand-collecting methods in any one cave. Gates (1959) probably encountered similar results since the mean number per cave based on the 173 earthworms he examined from 23 caves was 7.5. Although other methods of collection, in con- junction with the hand-sorting technique used in this study, might have revealed greater numbers in some cases, they would have caused great disruption to the cave ecosystem. The cave/ mine lengths examined ranged from 14 mto 122 m. Relative to the area of each cave, only a small amount of detritus is washed in. The amount of detritus rather than the length of the cave or mine appears to have a greater bearing on the number of earthworms at each site. Table 1 shows that although Harbells Cave is much longer than Glebe Pot, many more earth- worms were encountered in the latter. A sizeable stream runs through Harbells Cave and little detritus collects in this cave. Glebe Pot, on the other hand, collects and retains a great deal of detritus and large numbers of fallen leaves, both potential habitat and food material for earth- worms. Greenhead Cave is also longer than Glebe Pot. The entrance to Greenhead Cave, however, is on a cliff face and the exit is blocked by ceiling collapse. Virtually no detritus enters this cave and no earthworms were discovered though the cave was carefully searched. The source of energy and habitat material in Bat Cave is principally porcupine droppings, which are not favored by earthworms. No other significant amount of detritus appears to enter this cave, but more diligent searching may reveal THE CANADIAN FIELD-NATURALIST Vol. 91 362 sunp surdnoiod ‘snq0q snyijap pnur “‘JOARID) UMOPYeIIG YOO[q Jopuy [OAPIZ 19M (1 ‘314 22s) awop jo sulfIaD sso] JopuQ sdo] Jopuy) Jaquievyo jo [jen Ul jay90d [ews Ul snqiIEq pnu ul yoo1 JapupQ d0UI]UD JO [[eM UO SUI[MeID }e9S YUIW JIopuy) [}os uy sunp suidno10g yOOI Japun jos uy] Iaquiyy UIW UdT[e} UT yORID JOquil] 9UIW Jopuy) pnul ul x01 Japuy snjqijap uy BO] Ud}}01 Jopu_ pnw ul 4901 Jopuy yeqeH AP WI AP WI xP xP AP xP xP yi xP AP xP xP xP 4P xP xP Ui AP u0Z (Jp) 2Uu0Z Y1ep JO (Yi) 2UOZ PjOYsosY} UI SOAvD YOIMSUNIG MAN Ul SUTIOMYIIVI 1OJ BBP UOTJDaT[OD puw salsadS—] A1aV_L QA Seo wont ENE GNIEGN = GNI GN SGN GN ert suowioads jo Joquinyy Snpigns snjiapospuag Dasod DIuUasIy Snpiqns snpiapospuag DIDjINIAaGN] DapojJasiodp DIDjINIAIGN] DapOjJaIodp DIDjINIA4aqGN] DapOJalIOdp ‘ds papojoassodpy ‘ds papoljassodp ‘ds papojoassiodp ‘ds papojoassodpy Snpiqna snji4pospuad S1A]SAd9] SNILQUINT DIDjINI4aqN] DapOJIa44ody snpigna snjipospuag DIDjJNI4aqN] DapOjJad4Odp S141SALI9] SNIUQUNT saisads W1OMYIeY 9L61 “AON FI 9L61 “AON F1 9L61 ‘3NV €Z 9L61 3NY ZZ LL6I “URL 6 9161 0 OL 9461 90 OL SL6I “IEW SI 9L61 390 €Z 9L61 90 €Z 9L61 “INL SZ 9L6I “JBW 9 9L61 290 LI 9L61 390 LI 9L61 38NV 8z 9L61 3NV 8Z 9161 90 € 9L61 290 € 9L61 3428 61 9161 342g 81 a1eq cv= poAsainsun Sv= peaAdAinsun paAdAinsun 09= peAadarnsun 001 CCl vl (wr) yisu9] asessed ule] 0D Heqiv “QARD 1eg ‘OD uyor jules “QABRD) PRIyUIIIH ‘O—D uyor jules ‘QAR SOMO ‘O—- uYor jules ‘QABD ST[9qIeH ‘O- ssuly ‘QABD SITY ‘O—= sdulry ‘OUT, 2991D ‘O*—D ssuly 10d 29319 UOIBIO] ABD 1977 earthworms at this site. It is worth noting that McAlpine found Bat Cave low in both abun- dance and diversity of all species of invertebrate fauna. Table | also shows that most of the worms were concentrated in the dark zone. The pro- bable reason is that most of the detritus seems to collect here. Earthworms were collected in the threshold zone only in Glebe Pot where water runs over the entrance lip and collects the detritus at the bottom of the shaft, and in Harbells Cave, which has a muddy threshold zone that merges with the forest floor. The dark zone offers the most stable environ- ment in the cave. The soils and detritus of the threshold zone, when present, are usually shallow and offer earthworms little protection from damaging sunlight. The dark zone, how- ever, has the coldest mean annual temperature of any part of the cave, and this is not animmediate asset to earthworm reproduction or develop- ment of troglophilic populations. The porcupine, Erethizon dorsatum, com- monly dens in many caves in the Maritimes. Its large dung piles are a major source of energy and support a varied invertebrate community (Calder and Bleakney 1965) in many caves in this region. Of the hypogean environments exam- ined, Glebe Mine and Bat Cave served as porcupine denning sites. Calder and Bleakney (1965) felt that the acidity of cave detritus, principally porcupine dung, was a factor limiting the presence of earthworms in Frenchman’s Cave. Samples of porcupine dung in French- man’s Cave ranged froma pH (1:1 H,O) of 5.1 in poorly decomposed detritus to 6.3 in thoroughly decomposed feces. Although the present investigations revealed only one immature Aporrectodea sp. from dung samples, work on pH tolerance among earth- worms (Bodenheimer 1935; Wherry 1924) would indicate considerable variation in acidity toler- ance between species. Reynolds (1977) states that D. rubidus seems quite acid tolerant. He notes that A. ruberculata has been recorded from soils of pH 4.8-7.5, Eisenia rosea (Savigny) from soils of pH 4.9-8.0, and Lumbricus terrestris L. from soils of pH 4.0-8.1. It does not seem that soil reaction per se is the factor limiting earthworm coloni- zation of caves. MCALPINE AND REYNOLDS: OLIGOCHAETA ECOLOGY, NEW BRUNSWICK CAVES 363 Manure piles harbor, from time totime, all the species collected in this study (Reynolds 1977). These feces, however, are composed primarily of partially digested plant material that the earth- worms normally feed on. Porcupines, in con- trast, feed largely on the cambium layer of various trees and on conifers in particular. Reynolds and Jordan (1975) rank conifers lowest on the earthworm palatability scale. Porcupine dung could then be considered close to unpalatable for earthworms. This may be a better explanation of why these feces appear to be a marginal habitat for the oligochaetes en- countered in caves and mines in New Brunswick. Stone and Ogles (1953) noted that high con- centrations of the mite Uropoda agitans may limit earthworm populations. Calder and Bleakney (1965) found that poorly decomposed porcupine dung supported high populations of Acarina, averaging 67.3/100 cc of this detritus. They did not record the above mite species, but their collection was not completely identified (Calder and Bleakney 1967). Further collections of mites from caves in Nova Scotia and New Brunswick by Moseley (1974, 1975) have not revealed this species. In the cave environment earthworms are certainly not exposed to a wide variety of predators as are epigean forms. McLeod (1954) reported the staphylinid beetle Quedius meso- melinus as a predator of earthworms. This insect has been collected in Kitts Cave (Moseley 1975), and McAlpine has noted a staphylinid, possibly this species, in Glebe Mine. It appears that the smoky shrew, Sorex fumeus, occasionally enters caves in this area (McAlpine 1976) and on one trip to Glebe Mine McAlpine observed a deer mouse, Peromyscus maniculatus, in the dark zone. Although Hamilton (1941) has found that earthworms make up a relatively insignificant part of the diets of these two mammals, they likely consume any earthworms they discover while foraging in caves or mines. Earthworms are probably commonly intro- duced with seasonal runoff into those caves that get a seasonal influx of detritus after heavy rains in spring or autumn. Gates’ (1961) division of megadrile species, based on their food habits, is of interest. He considered D. rubidus a litter feeder. He termed the other three species 364 ’ q Ree ee PS 4 ws” x. THE CANADIAN FIELD-NATURALIST (ees 1k ‘' oe ia on : A eas aN Be By 6 Vol. 91 FIGURE 1. Dendrodrilus rubidus crawling on the ceiling dome of Howes Cave, New Brunswick. collected in this study geophagous (those species that pass much soil through their intestines). If such is the case, D. rubidus would be more commonly at the soil surface and most prone to be washed into caves. Earthworms may also enter caves or mines under their own power. Dendrodrilus rubidus seems to be an active species. Reynolds (1977) reported that on damp nights it has been noted crawling on the surface of the ground and even climbing trees. In Glebe Mine, earthworm cocoons and castings, probably of D. rubidus, were observed on ceiling beams. In Howes Cave a specimen of D. rubidus was collected from the ceiling of a small dome (see Figure 1). A single specimen of A. tuberculata was noted crawling about 20 cm above the ground on the damp wall of the threshold zone in the entrance to Harbells Cave. This same species was collected in the dark zone of this cave. In Glebe Mine, which does not receive the same seasonal influx of plant detritus with runoff as do the caves, active colonization may be of some importance in earthworm introduction. In the particular case of mines man may play as important a role in introducing earthworms to the habitat as do seasonal weather patterns. In Glebe Mine, the only food available to earth- worms in any great quantity in the dark zone are support timbers, both standing and fallen. If these had been stored outside for a while, it is not difficult to understand how earthworms or their cocoons could have been transported into the mine with clumps of soil adhering to the beams. The mine was opened about 100 years ago. Voisin (1961) found the optimum soil temper- ature fora number of earthworm species, two of which were collected in this study. Optimum soil temperatures for D. rubidus are 18-20° Cand for E. rosea, 12°C. Interestingly, these two worms represent the extremes of the optimum soil temperatures for the seven species he examined. Bodenheimer (1935), working with A/dlolobo- phora samarigera Rosa, found the range of normal activity to be 14.7-27.8°C. He found only “interrupted crawling” at 14.7°C, “only weak movements” at 6.7°C, and the “beginning of cold torpor” at -1.3°C. Reynolds (1977) and Gates (1961) have noted that earthworms may be active under ice and snow or in litter surrounded by snow. Specimens of D. rubidus have been collected in breeding condition from a ridge of soil surrounded by 75 cm of snow. This cold weather activity would 1977 indicate that local populations may at least be pre-adapted enough to remain normally active in caves and mines when introduced into these environments. The work by Voisin (1961) would indicate that some study is needed to determine whether all megadrile species collected in this investigation are able to reproduce in Maritime caves and mines without a period of acclimati- zation. Reynolds (1977) noted that estivation and hibernation are climatically imposed in Ontario for the four species collected in this study, with reproduction being limited to autumn and spring. McAlpine has noted ice in the outer dark zone of some New Brunswick caves in July. Brief observations on air temperatures in caves in southern New Brunswick by McAlpine and more detailed work by Calder and Bleakney (1965) in Nova Scotia have revealed that cave air temperatures in the constant-temperature zone hover around 3-4°C in both regions. It is generally known that cave air temperatures in this zone approximate the mean annual surface temperatures for the areas in which they are situated. Due to the short length of many New Brunswick caves constant temperature zones may not be present in all caves. Calder and Bleakney (1965) found temperature readings to be 1°C cooler 5 cm below the surface of detritus than in the adjacent air. Although some acclimatization may take place, a temperature of 4°C must affect the reproductive and metabolic rates of earthworms in caves, and under these temperature conditions it would appear that earthworms would neither reproduce quickly nor require large food sup- plies of detritus. Dendrodrilus rubidus was very often asso- ciated with branches on the cave floor or mine support beams. Reynolds frequently encounters this species in northeastern North America under the bark of decaying logs. Dendrodrilus rubidus was collected a number of times from within cracks in fallen beams. When associated with timbers, A. tuberculata was beneath them in the detritus or shallow soils. Usually A. tuber- culata was collected from pockets of detritus or in mud under rocks. It was interesting to note that in Kitts Cave, where beaver (Castor cana- densis) have constructed dams and stored caches MCALPINE AND REYNOLDS: OLIGOCHAETA ECOLOGY, NEW BRUNSWICK CAVES 365 of Salix and Alnus (McAlpine 1977), two favored earthworm foods (Reynolds and Jordan 1975), D. rubidus was associated with these beaver caches. Conclusions The origin of cavernicolous megadrile popula- tions are commonly accidental, through the actions of weather or man, but may be through active colonization. The structure of the en- trance to the cave or mine probably controls whether passive or active modes of colonization are of prime importance in the introduction of earthworms to any given cave or mine. Once the initial introduction has been made, reproduction may or may not take place. In New Brunswick, annual seasonal introductions of individuals with detritus may ensure the presence of earth- worms within the cave from year to year even though the earthworms may not be reproducing in the caves. These earthworms could be con- sidered trogloxenic. Evidence presented earlier that D. rubidus is reproducing within Glebe Mine would indicate that some individuals of this species are troglophilic. The immature A. tuberculata collected from dung samples in the mine lend support to the conclusion that this species may also be reproducing in Glebe Mine. It is also worth considering whether earth- worm reproduction in New Brunswick caves is limited to the variable-temperature zone ini- tially. Although this cave zone is much cooler during the summer months than are nearby epigean earthworm habitats, its mean annual high temperature is above that of the constant- temperature zone. Acclimatization may still be necessary, but the demanded reproductive adaptation to temperature would not be as severe as in the deeper and colder parts of the cave Or mine. Gates (1959) has reported D. rubidus to be the common cave earthworm in the United States. It was the only species he reported from a mine in that country and it has been collected in mines in Canada and France as well. Dendrodrilus rubidus is a litter form, prone to be washed into caves, is commonly associated with logs (Rey- nolds 1977) and mine support beams, is easily introduced to caves and mines by man, and appears to crawl actively some distances over the ground surface. These may be reasons to explain 366 why it is socommonly encountered in both caves and mines. In Britain Eiseniella tetraedra (Savigny) is the most common earthworm in caves (Reynolds 1977). Although the species has been recorded in New Brunswick (Reynolds 1976b), it has not been collected in any of the counties where caves in this study are located, or in any Canadian caves. Based on the information presented here and the limited work of others on the topic, D. rubidus seems to be the common Canadian cave earthworm. Acknowledgments We thank the following for their help in the field: Glendon MacDonald, Roy McAlpine, Thomas McAlpine, Max Moseley, Cynthia Regan, and Bill Walton. We also acknowledge Bill Staples for producing the figure in the text. We are grateful to David Christie and Max Moseley who read an early draft of the manu- script and provided comments that led to its improvement. Literature Cited Bodenheimer, F.S. 1935. Soil conditions which limit earthworm distribution. Zoogeographica 2: 572-578. Calder, D. R. and J.S. Bleakney. 1965. Microarthropod ecology of a porcupine inhabited cave in Nova Scotia. Ecology 46: 895-899. Calder, D. R. and J.S. Bleakney. 1967. Observations on Frenchman’s Cave, Nova Scotia and its fauna. Bulletin of the National Speleological Society 29(1): 23-25. Cook, D.G. 1975. Cave-dwelling aquatic Oligochaeta (Annelida) from the eastern United States. Transactions of the American Microscopical Society 94(1): 24-37. Gates, G. E. 1959. Earthworms of North American caves. Bulletin of the National Speleological Society 21(2): 77-84. Gates, G.E. 1961. Ecology of some earthworms with special reference to seasonal activity. American Midland Naturalist 66(1): 61-86. Gates, G. E. 1970. Miscellanea Megadrilogica VIII. Miega- drilogica 1(2): 1-14. Gates, G. FE. 1972. Toward a revision of the earthworm family Lumbricidae. IV. The trapezoides species group. Bulletin of the Tall Timbers Research Station Number 12. 146 pp. Halliday, W. R. and D.I. Dunnet. 1966. Reconnaissance of a new rain-forest karst area on Vancouver Island, British Columbia, Canada. Vancouver Island Speleo- logical Survey Bulletin 2, Western Speleological Survey Bulletin 38, August 1966. Reprinted in Speleological THE CANADIAN FIELD-NATURALIST Vol. 91 Digest 1966. pp. 2-40 — 2-47. Hamilton, W. J. 1941. The food of small forest mammals in the eastern United States. Journal of Mammalogy 22: 250-263. McAlpine, D.F. 1976. Howe’s Cave, New Brunswick. Canadian Caver 8: 28-30. McAlpine, D. F. 1977. Notes on cave utilization by beaver. Bulletin of the National Speleological Society 39(3): 90-91. McLeod, J. H. 1954. Notes on a staphylinid (Coleoptera) predator of earthworms. Canadian Entomologist 86(5): 236. Moseley, C. M. 1974. N.S.S.S. biological records. Part 1. Nova Scotia Speleological Society Newsletter 3: 9-11. Moseley, C. M. 1975. N.S.S.S. biological records. Num- ber 2. Nova Scotia Speleological Society Newsletter 6: 8-11. Nicholas, G. 1960. Checklist of macroscopic troglobitic organisms of the United States. American Midland Naturalist 64(2): 123-160. Peck, S. B. and M. B. Fenton. 1973. The fauna of Cana- dian caves. Canadian Caver 5: 18-23. Reynolds, J. W. 1974. Are oligochaetes really herma- phroditic amphimictic organisms? Biologist 56(2): 90-99. Reynolds, J. W. 1975. Die Biogeografie van Noorde- Amerikaanse Erdwurms (Oligochaeta) noorde van Meksiko — I. Indikator 7(4): 11-20. Reynolds, J. W. 1976a. Die Biogeografie van Noorde- Amerikaanse Erdwurms (Oligochaeta) noorde van Meksiko — II. Indikator 8(1): 6-20. Reynolds, J. W. 1976b. A preliminary checklist and distri- bution of the earthworms of New Brunswick. New Bruns- wick Naturalist 7(2): 16-17. Reynolds, J. W. 1977. The earthworms (Lumbricidae and Sparganophilidae) of Ontario. Royal Ontario Museum Life Sciences Miscellaneous Publications. x + 141 pp. Reynolds, J. W. and D.S. Christie. 1977. Additional earthworm records for New Brunswick. New Brunswick Naturalist 8(3): 25. Reynolds, J. W. and G.A. Jordan. 1975. A preliminary conceptual model of megadrile activity and abundance in the Haliburton Highlands. Megadrilogica 2(2): 1-11. Stone, P. C. and G. D. Ogles. 1953. Uropoga agitans, a mite pest in commercial fishworm beds. Journal of Economic Entomology 46(4): 711. Vandel, A. 1965. Biospeleology. Pergamon Press, London. 524 pp. Voisin, A. 1961. Lazy and active earthworms. /n The challenge of earthworm research. Edited by R. Rodale. Soil and Health Foundation, Emmaus, Pennsylvania. pp. 9-13. Wherry, E. T. 1924. Soil acidity preference in earthworms. Ecology 5(3): 309. Zicsi, A. 1974. Ein neuer Hohlen-Regenwurm (Oligochaeta: Lumbricidae) aus Ungarn. Acta Zoologica Academiae Scientiarum Hungaricae 20(1-2): 227-232. Received 11 March 1977 Accepted 23 July 1977 Summer Habitat Use by White-tailed Ptarmigan in Southwestern Alberta PATRICK W. HERZOG Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9 Herzog, Patrick W. 1977. Summer habitat use by White-tailed Ptarmigan in southwestern Alberta. Canadian Field- Naturalist 91(4): 367-371. Periodic censuses of White-tailed Ptarmigan (Lagopus leucurus) in alpine basins of southwestern Alberta between 15 July and 31 August 1976, provided information on habitats used during the summer season. Birds occupied areas in which rocks covered about half of the ground surface and where water was available within 25 m. Willows (Salix spp.), mosses (Bryophyta), grasses (Poaceae), saxifrage (Saxifraga spp.), sedges (Carex spp.), groundsel (Senecio spp.), cinquefoil (Potentilla spp.), and everlasting (Antennaria spp.) were the major components of vegetation in areas used by ptarmigan. Males and broodless females moved away from breeding ranges in late July; females with broods remained on the breeding range but frequently travelled downhill along stream courses even below treeline. The cause of this apparent segregation and its extent are unknown. Key Words: White-tailed Ptarmigan, summer habitat, Alberta. White-tailed Ptarmigan occupy alpine areas with an abundance of rock and low-growing vegetation during the summer season (Weeden 1959; Choate 1963; Braun 1971). In Colorado most males and broodless females move from breeding areas at lower limits of the alpine tundra upslope in early July to higher ridges where late-lying snowfields provide sufficient moisture to support vegetation used as food (Braun 1971). Females with broods also move upslope during the summer, but generally remain at somewhat lower elevations; sedges (Carex spp.), avens (Geum spp.), and clovers (Trifolium spp.) are common plants in the areas used by all birds (Braun 1971). Choate (1963) also observed ptarmigan following the retreating snow upslope during the summer months. He recorded snow willow (Salix nivalis), heaths (Cassiope spp. and Phyllodoce spp.), and mosses as the dominant plants at sites where ptarmigan were observed from early June to mid-Sep- tember. These study areas in Colorado and Montana are characterized by large expanses of gently rolling alpine tundra (Choate 1963; Braun and Rogers 1971). Conversely, in southwestern Alberta, White-tailed Ptarmigan breed in small alpine basins above which relief is steep and vegetation immediately upslope rapidly be- comes sparse. Terraces of alpine tundra, which might provide late-lying snowfields, do not exist. Habitat use varies with geologic situations, thus this study was undertaken to determine the vegetative components of ptarmigan summer habitat in southwestern Alberta and whether the sexes differed in their use of habitat. Study Areas Ptarmigan were censused in the Rocky Mountains Forest Reserve approximately 9-10 km east of Kananaskis Lakes and 42 km west of Turner Valley, Alberta, at elevations of 2170-2640 m. Three areas were chosen for study: Burns Lake (50°36’ N, 114°57’ W), High- wood Pass (50° 36’ N, 114°59’ W), and the basin between Elpoca and Tombstone Mountains locally known as Goat Valley (50°40’N, 115°02’ W). These areas provide a mixture of krumholz, rock slides, cliffs, willow shrubs, and large herbaceous mats of heather (Cassiope tetragona), heath (Phyllodoce glanduliflora), and mountain avens (Dryas hookeriana). Streams, which flow throughout the summer, drain these basins; snowfields rarely persist past mid-July. Methods Study areas were searched for ptarmigan at intervals of 7 to 10 days from 15 July to 31 August 1976. Birds were located by systemati- cally searching all cover with the aid of pointing dogs. Some bias in the census technique occur- red, as an attempt was made to capture all un- marked birds encountered; thus more time was spent in areas where birds previously had been located. I believe this bias is small, because all areas were searched thoroughly, and a special effort was made in areas where ptarmigan sign 367 368 THE CANADIAN FIELD-NATURALIST Vol. 91 TABLE 1—Percent cover by rocks, water, and various species of plants in habitat used by White-tailed Ptarmigan during the summer season (15 to 31 August 1976). Data from 82 census plots. Others include all species with frequencies less than 15% Vegetation Rocks Water Mosses (Bryophyta) Grasses (Poaceae) Saxifrage (Saxifraga lyalli, S. bronchialis, and S. oppositifolia) Snow Willow (Salix nivalis) Sedge (Carex spp.) Groundsel (Senecio lugens and S. triangularis) Cinquefoil (Pontentilla diversifolia and P. fruticosa) Arctic Willow (Salix arctica) Everlasting (Antennaria alpina var. media and A. lanata) Common Yarrow (Achillea millefolium) Drummond’s Anemone (Anemone drummondi) Moss Campion (Silene acaulis) Flea-bane (Erigeron aureus, E. humulis, and E. grandiflorus) Fireweed (Epilobium alpinum and E. latifolium) Alpine Speedwell (Veronica alpina) Grass-of-Parnassus (Parnassia montanensis) Mountain Sorrel (Oxyria digyna) Indian Paintbrush (Castilleja occidentalis) Alpine Forget-me-not (Myosotis alpestris) Smelowskia calycina Heart-leaved Arnica (Arnica cordifolia) Mountain Avens (Dryas hookeriana) Rock Willow (Salix vestita) Milk Vetch (Astragalus occidentalis) Saussurea densa Snow Buttercup (Ranunculus eschscholtzii) Yellow Mountain Heath (Phyllodoce glanduliflora) Others [Elephant Head (Pedicularis groenlandica), Stonecrop (Sedum rosea and S. stenopetalum), Blue-green Gentian (Gentiana glauca), Sandwort (Arenaria sajanensis), Dwarf Alpine Dandelion (Taraxacum lyratum), Rocky Mountain Goldenrod (Solidago multiradiata), Sweet-flowered Androsace (Androsace chamaejasme), Slender Blue Beard-tongue (Penstemon procerus), Hedysarum sulphurescens, White Camas (Zygadenus elegans), Rocky Mountain White Heather (Cassiope tetragona), Wild Strawberry (Fragaria virginiana), Low Larkspur (Delphinium bicolor), and Horsetail (Equisetum scirpoides)] (feathers and/or droppings) was found but birds were not observed. A circular plot of 50 m?2(ca. 4.0 m radius) was centered on the point of location where an individual bird was first sighted. The location of the female was chosen as plot center for females with broods. The presence and relative abun- dance of all species of plants within the plot were determined. Plants were identified using the flora by Moss (1959), and the percent coverage Frequency of occurrence (%) Average cover (%) 100 45 78 73 65 61 pS m= = NUON UON == KE DNKH WH NH NNO—-WwWNOORWOD O — ise) in — nN — | of individual species was visually estimated to the nearest 5% (2.5 m2). Species comprising less than 5% coverage of the plot were recorded as being present and were assigned a value of 1% in quantitative summaries. The distance of ptarmigan locations to the nearest willow shrub greater than 30cm in height, coniferous cover, and source of water (snowfield, stream, or lake) was recorded to determine the extent to which these habitat 1977 HERZOG: WHITE-TAILED PTARMIGAN, ALBERTA 369 TABLE 2—Number of sightings (%)of White-tailed Ptarmigan within 25 m of nearest water source, willow cover greater than 30 cm in height, coniferous cover, and the mean distance (m) + SD to these habitat features. Sightings of females with broods (N=48) and without broods (N =7) are summarized separately No. of sightings < 25 m (%) Mean distance (m) + 1 SD Habitat Males Females Females Males Females Females feature (N = 27) (N = 48) (N = 7) Total (N = 27) (N = 48) (N = 7) Average Water 19(70) 42(88) 6(85) 67(82) 8x6 7/2833} 10+6 sta Willow 7(26) 29(60) 0(0) 36(44) 1111 yas 7/ = s"7/ Conifer 4(15) 13(27) 0(0) 17(21) 8t7 1649 = 14+8 features might influence ptarmigan distribution. Distances up to 100 m were recorded to the nearest meter; habitat features beyond 100 m were recorded as being distant and were not included in quantitative analysis. Results and Discussion Ptarmigan were seldom located far from boulders and/or rock slides; rocks occurred with both the highest frequency (100%) and percent coverage (49%) in the census plots (Table 1). Rocks provide essential cover for these ptar- migan, and birds are usually observed in areas where rocks are greater than 30 cm (12 in) in diameter (Weeden 1959; Choate 1963; Braun 1971). Similarly, with the exception of males located late in summer, ptarmigan were usually located within 25 m of a snowfield, stream, or lake (82% of all sightings) (Table 2). Thus, ptarmigan were concentrated along stream courses with an abundance of rocks (Figure 1) and were rarely observed in areas that did not provide both rocks and moisture (e.g., fore- ground, Figure 2). Willow shrubs and conifers (Picea spp., Abies spp., and Pinus spp.) greater than 30 m (12 in) in height were considered as alternate sources of potential cover (Table 2). As they travelled along streambanks females with broods were often in close proximity to willow shrubs (mean distance of 12m, 60% of all sightings), but birds did not crouch in this vegetation when disturbed. Ptarmigan preferred to escape by running and/ or flying, or to remain motionless near rocks at the water’s edge. No group of birds was associated with coniferous cover to any degree (Table 2). These results confirm the observations of Choate (1963) that ptarmigan are rarely found in vegetation taller than themselves, and thus are dependent almost solely on rocks for cover presumably from inclement weather and predators. Rocks also trap drifting snow, which might provide mois- ture for plants as the summer progresses. The presence and abundance of species of vegetation in habitat used by ptarmigan are presented in Table |. Willows, grasses, sedges, cinquefoil, and everlasting were main com- ponents of the vegetation; mosses, saxifrage, and groundsel were also very common, and reflect the moist conditions where ptarmigan were found (Table 1). In contrast to Choate’s (1963) study area, heath and heather were. minor components of the vegetation at my sites. Weeden (1959) also rarely observed White-tailed Ptarmigan in such vegetation, and although large mats of heath, heather, and mountain avens were present in my study areas, birds avoided these areas (e.g., foreground, Figure 2). These sites were typically open with few boulders and were very dry early in summer. Weeden (1959) also thought broods might have dif- ficulty travelling through dense vegetation (heath and heather) and so avoided these areas. Several species of plants were present regularly (Table 1) but contributed little to the total coverage of census plots owing to their nature of growth and relative abundance. Vegetation at census plots was similar for male and female ptarmigan, and results were combined in Table |. But 18 of the 27 sightings of males (67%) occurred prior to 7 August when males were still occupying breeding range. After 7 August, males were located on only nine occasions; four of these sightings were high on windswept ridges where vegetation was sparse (Figure 2) and the other five occurred on 29 and 30 August near to the July areas. Only seven 370 ieee FIGURE 1. Streamside habitat used by all White-tailed Ptarmigan in early summer and by females with broods throughout the summer. THE CANADIAN FIELD-NATURALIST Vol. 91 sightings were obtained of females without broods and these all occurred prior to 1 August. More observations of males and broodless females are needed in mid- and late summer to determine whether there are significant dif- ferences between the type of vegetation used by these groups and females with broods. Movement away from spring territories by male and broodless female ptarmigan to higher elevations has been reported in Montana and Colorado (Choate 1963; Braun 1971). Obser- vations of males high on ridges in August, the failure to locate any males in the Goat Valley study area after 3 August, and the inability to locate leg-banded individuals other than females with broods after 7 August provide evidence for summer movements away from breeding areas in southwestern Alberta. Extensive movements to higher elevations above breeding areas may not be possible because of the geologic structure of the Rocky Mountains in this part of the bird’s range; individuals may move to distant basins at higher altitudes. The summer movements of males and broodless females is an intriguing problem for future research. Females with broods occupied the same areas throughout the summer and concentrated their activities along streambanks (Table 2). Unlike the situation elsewhere of females with broods gradually moving upslope to higher elevations (Choate 1963; Braun 1971), females with broods were located on the high ridge in the background on three occasions in mid-summer. 1977 in this study frequently moved downslope along stream courses (often below tree-line). In areas intensively grazed by livestock, horizontal and downhill movement by both sexes of ptarmigan can occur (Braun 1971). The presence of certain species of plants in census plots used by ptarmigan may reflect food habits to some degree. Weeden (1967) found the leaves of willows and buttercups (Ranunculus spp.), seeds of sedges and grasses, and fruits of bistorts (Polygonum spp.) to be common items in the diet of adult ptarmigan in summer. May and Braun (1972) agreed with these findings and indicated that females selectively fed on bistorts while males ate the seeds and leaves of plants according to their availability, including clovers (Trifolium spp.), mouse-ear (Cerastium spp.), mustards (Cruciferae), snowball saxifrage (Saxi- fraga rhomboidea), and alpine avens (Geum rossi). Bistorts, clovers, and alpine avens were not present in my study areas and alternate food items are probably selected; females with broods were observed feeding repeatedly on the flowers and leaves of Smelowskia calycina (Cruciferae), a food item not previously recorded (Weeden 1967; May and Braun 1972). As suggested by May and Braun (1972), ptarmigan distribution in summer may be closely related to the availability of green vegetation, and indeed differences in food habits between the sexes may influence habitat selection. In summary, alpine areas with an abundance of rocks for cover, and where sufficient moisture for the growth of fresh vegetation is available, are used during the summer by White-tailed Ptarmigan in southwestern Alberta. All birds use these spring breeding areas to meet habitat requirements early in summer; females with broods continue to use these areas throughout the summer. I believe this use constitutes selection because wide expanses of potential habitat are avoided. A definite segregation of HERZOG: WHITE-TAILED PTARMIGAN, ALBERTA Sil habitat type occurs in mid-summer: males and females without broods move away from breeding basins (possibly to distant basins of higher elevation), but females with broods remain along stream courses, frequently travel- ling downstream below tree-line. The extent of seasonal movements is unknown, and the reason different members of the sexes segregate into different habitat is not apparent. Acknowledgments I thank Madeleine Dumais, Assistant Curator of the Herbarium, Department of Botany, University of Alberta, for aid in identification of plant species; and D. A. Boag, Department of Zoology, University of Alberta, for reviewing the manuscript and offering many helpful suggestions, and helping to defray the costs of publication. Literature Cited Braun, C.E. 1971. Habitat requirements of Colorado White-tailed Ptarmigan. Proceedings of the Western Association of State Game and Fish Commissioners 51: 284-292. Braun, C.E. and G.E. Rogers. 1971. The White-tailed Ptarmigan in Colorado. Colorado Division of Game, Fish and Parks Technical Publication 27. 80 pp. Choate, T.S. 1963. Habitat and population dynamics of White-tailed Ptarmigan in Montana. Journal of Wild- life Management 27(4): 684-699. May, T. A.and C. E. Braun. 1972. Seasonal foods of adult White-tailed Ptarmigan in Colorado. Journal of Wild- life Management 36(4): 1180-1186. Moss, E. H. 1959. Flora of Alberta. University of Toronto Press, Toronto. 546 pp. Weeden, R.B. 1959. The ecology and distribution of ptarmigan in western North America. Ph.D. thesis, University of British Columbia, Vancouver. 247 pp. Weeden, R. B. 1967. Seasonal and geographic variation in the foods of adult White-tailed Ptarmigan. Condor 69(3): 303-309. Received 17 February 1977 Accepted 25 July 1977 Horned Grebe Breeding Habitat in Saskatchewan Parklands LAWSON G. SUGDEN Canadian Wildlife Service, Saskatoon, Saskatchewan S7N 0X4 Sugden, Lawson G. 1977. Horned Grebe breeding habitat in Saskatchewan parklands. Canadian Field-Naturalist 91(4): 372-376. Distribution of Horned Grebes, Podiceps auritus, on a 31.1-km? area with 25 ponds per km? was measured in 1974 and 1975. Before, and particularly during, the nesting season the grebes tended to select permanent ponds over 0.2 ha that had areas without emergent vegetation. Distribution was not affected by tree growth around ponds, but prior to nesting the birds showed some preference for ponds surrounded by pasture rather than cultivated fields. There were 47 nests on 47 ponds (1.5 nests per km?) in 1974, and 103 nests on 102 ponds (3.3 nests per km?) in 1975. Key Words: Horned Grebe, Podiceps auritus, breeding habitat, nesting, Saskatchewan, parklands. Habitat used by breeding Horned Grebes (Podiceps auritus) has been studied in detail in Norway, Iceland, and Finland by Fjeldsa (1973a, b), and has been described fora grassland area in North Dakota (Faaborg 1976), but quantitative descriptions are lacking for parkland habitat where, in North America, the species apparently reaches its highest densities. With increasing pressure on wildlife habitat from other land uses, data on habitat requirements should be gathered and documented for bird species that may be threatened by loss of habitat. This study documents nest densities and describes the habitat used by Horned Grebes ona study area in Saskatchewan during 1974 and 1975. Study Area The study area (52°N, 106°W) is a block, 4.83 X 6.44 km, 48 km east of Saskatoon. It is divided by roads and fencelines into 12 sections, each 2.59 km? (1 mi2), and these were used as sampling units. The block comprises Sections 14-17, 20-23, and 26-29 of Township 36, Range 28, West 2. The area is in the aspen parkland (Bird 1961), soils are dark brown, and topog- raphy is rolling to gently rolling (Mitchell et al. 1947). Annual precipitation averages about 36cm. Land use on the study area was exemplified by 1975 percentages: grain (barley and wheat), 41.5; summer fallow, 29.4; rapeseed, 5.9; pasture, 6.1, about one-half of which was grazed; farmyards and roads, 1.2; wasteland (chiefly uncultivated areas around ponds), 6.3; and ponds, 9.6. Pond density averaged 25/km2. Size ranged from less than 0.04 ha to 8.1 ha. Most ponds were partly or wholly bordered with trees, mainly willows (Salix spp.) and trembling aspen (Populus tremuloides). Limnological data for the ponds are lacking; however, these ponds are similar to the numerous closed, eutrophic, freshwater ponds found throughout the prairie- parkland region (Driver 1977; Driver and Peden 1977). The wetland vegetation on the area followed closely the lists given by Millar (1976, p. 22) for species found in various freshwater vegetation zones of prairie wetlands. One exception was the scarcity of Scirpus spp. on my area. The most common emergent species occurring in ponds used by grebes were Typha latifolia, Scolochloa festucacea, and Carex atherodes. Methods Maximum depth of ponds was measured in early May and again in July. Ponds were assigned to permanency categories similar to those described by Martin et al. (1953) and Evans and Black (1956). Type | ponds are shallow depressions, seldom having over 25 cm of water and, except in wet years, dry up before July. Type 3 ponds are shallow marshes usually not more than 60 cm deep. These ponds also tend to dry up during summer except in wet years. Type 1 and 3 ponds that contain water through the growing season usually become overgrown with emergent vegetation. Type 4 ponds have up to 120 cm of water in the spring and seldom become dry. Growth of emergent vegetation such as Scolochloa and Carex leaves little or no open water by July. Type 5 ponds are over 120 cm 372 ST. deep in spring and contain water even in dry years. Emergent vegetation usually occurs only around the pond margin and a majority of the area remains open. This is the main distinction between Type 4 and Type 5 ponds. Pond areas were measured from maps prepared from aerial photographs. For data analysis, ponds were assigned to six area classes: < 0.21; 0.21-0.40; 0.41-0.81; 0.82-1.62; 1.63-3.24; 3.24 ha. Similarly, the percentage of shoreline bordered by trees (mainly Salix spp.) was measured and each pond was assigned to one of three cate- gories of woody shore growth: open, 0-33%; half-open, 34-66%; and closed, 67-100% (Smith 1971). Land use around ponds was re- corded in late May as summer fallow, seeded to grain, seeded to oilseed, or as pasture. Horned Grebes were censused three times in May each year. Each pond was visited in one day between 0830 and 1530 hours by one of four people making the census. From late May to late July, all ponds were searched twice for nests. Eggs in a nest, or, on some occasions, a brood on a pond were criteria used to establish the presence of a nest. The effect of the different pond variables on the distribution of birds and nests was tested with chi-square. Results and Discussion Some Habitat Features Figures for the numbers of ponds by per- manency type and size class in 1974 and 1975 were pooled (see Table 2). For each category, numbers in the two years were similar. Mean area of Type | ponds was 0.07 ha; Type 3, 0.15 ha; Type 4, 0.42 ha; and Type 5 ponds, 1.17 ha. Because Type | and 3 ponds received negligible use by grebes, they were excluded from further analyses. Numbers of Types 4 and 5 ponds per 259-ha section varied from about 13 to 58 in both years. The total numbers of Types 4 and 5 ponds were 455 and 453 in 1974 and 1975, respectively. Percentages of ponds in the three categories of peripheral tree growth were similar in both years: 43% open, 14% half-open, and 43% closed. Water levels were above average during the study and much of the peripheral tree growth was inundated throughout the nesting seasons. SUGDEN: HORNED GREBE BREEDING HABITAT, SASKATCHEWAN 3/8: Pre-nesting Distribution Numbers of ponds occupied and numbers of Horned Grebes seen during May counts in 1974 and 1975 are given in Table 1. Results of counts are comparable to the extent that the same technique was used throughout. Some birds were probably missed during counts, particularly on ponds with extensive flooded willows. At times Horned Grebes on open water swam into such cover when alarmed. Some of the single birds seen probably represented pairs. Also, nests were found on some ponds where no birds had been seen during counts suggesting that they had been missed, though movement onto such ponds after counts could not be ruled out. For these reasons the figures can be used only as population indices. Because more than one nest on a pond was a rarity, the number of ponds on which one or more birds was seen perhaps would be the best index to breeding pairs in the absence of data for nest densities. The first count in 1974 and the last in 1975 were chronologically similar and therefore were compared using a paired t-test for the 12 sections. There were no significant differences (P > 0.05) between years in either total birds seen or numbers of ponds occupied. Horned Grebes were never seen on Type 1 ponds. In 1974, they were recorded on four Type 3 ponds during counts, and in 1975, on one. The first year, 30% were seen on Type 4 ponds compared with 68% on Type 5 ponds. The 1975 figures were 22% and 77% for Types 4 and 5 ponds, respectively. In both years these propor- TABLE 1—Comparison of numbers of ponds occupied by various numbers of Horned Grebes on a 31.1-km? area during 1974 and 1975 counts Number of ponds Date in May 1974 Date in May 1975 Number of birds 17 23 24 8 12 15 l 18 24 24 14 1] 15 y) 51 44 34 17 29 39 3 0 1 0 0 I 1 4 3 0 l 0 0 0 5 0 0 0 | 0 0 6 0 0 I 0 0 0 Total number Ponds with birds 72 69 60 32 41 55 Birds IS NS NO 53 72 96 374 tions were significantly different (P< 0.001) from those expected if distribution had been proportional to the numbers of Types 4 and 5 ponds present. The grebes showed a strong preference for the permanent Type 5 ponds that contained patches of open water. Faaborg (1976) also found that Horned Grebes tended to select open ponds, but permanency type seemed unimportant in that study. The preference for Types 4 and 5 ponds was reflected in the grebe distribution on the study area. Numbers of birds on individual 259-ha sections were directly correlated with numbers of Types 4 and 5 ponds in both years (r = 0.8; P< 0.001). Size of ponds also influenced the distribution of Horned Grebes, with higher use shown for large ponds. This tendency likely accounted for part of the disproportionate use of Types 4 and 5 ponds because the average area of the latter was greater. In 1974, the distribution was sig- nificantly non-random (P< 0.001) with regard to size on both Type 4 and Type 5 ponds. Most of the difference was due to the low use of ponds under 0.2 ha and, conversely, high use of ponds over 0.4 ha. Use of Type 5 ponds in 1975 was likewise related to size (P < 0.001), but the small sample for Type 4 ponds did not reveal any significant difference (P > 0.05) with regard to size. The presence or absence of tree growth around ponds had no significant (P > 0.05) effect on the use of ponds by Horned Grebes in either year. They used all categories of ponds including those completely closed and less than 0.2 ha. Dwyer (1970) compared numbers of Horned Grebes on two Manitoba areas of equal size, one in a park with tree growth around all ponds and one 8 km from the first but on agricultural land where ponds were largely devoid of trees. Considerably more pairs were seen on the agricultural area and Dwyer (1970) concluded that tree growth around the ponds in the park deterred the birds. My results indicate that other factors probably were responsible for the observed differences in Horned Grebe numbers between the two Manitoba areas. In both years, Horned Grebes tended to use ponds surrounded by pasture in preference to those surrounded by cultivated land. Tests showed significance (P< 0.05) for all three counts in 1974, and the last two in 1975 THE CANADIAN FIELD-NATURALIST Vol. 91 (P< 0.02 and P< 0.01). Reasons for selection of pasture ponds are unknown, but may have involved an innate preference for a relatively pristine landscape, differences in food resources between the ponds, or disturbance by farming activities on cultivated land. Nesting Chronology Based on a laying rate of an egg every 2 days (DuBois 1919), and in some cases, size of young, most of the clutches in both years had been started in late May or early June. In 1974 at least 16 nests had been started in May and 38 in 1975. Perhaps the earliest were two nests, each with eight eggs in early incubation, found on 30 and 31 May. Four nests in 1974 and 18 in 1975 had been started in late June or early July, considerably later than average. These may have been re-nesting attempts because Horned Grebes readily re-nest if their first nest is destroyed (DuBois 1919; Fjeldsa 1973b). My data on nesting chronology contrast with those of Munro (1941) for a similar latitude in British Columbia, where it was unusual for eggs to be laid before the last week of June or early July. Egg dates given by Bent (1963), however, indicate that the Horned Grebes on my area did not nest unusually early. Distribution of Nests In 1974, 47 Horned Grebe nests or broods were found on the study area for a density of 1.5 km’. All were on different ponds. There were 103 or 3.3 per km’ in 1975. Reasons for the difference in nesting effort between the two years (see figures for adults in Table 1) are unknown. Evidently there were considerable numbers of non-breeding birds in the 1974 population, whereas most of the birds observed in 1975 were breeders. Munro (1941) stated that it was usual to find small numbers of non-breeding Horned Grebes on his British Columbia area. Fjeldsa (1973b) reported that 9 to 10% of the birds did not breed in Iceland and Norway. There, the rate of non-breeding decreased with increasing lake productivity (standing crop of benthos). Also, non-breeding birds apparently were mostly first- year birds unable to compete successfully for suitable territories. Nests were not found on Types | and 3 ponds (Table 2). Obviously, the temporary nature of these ponds in most years would create a lethal 1977 SUGDEN: HORNED GREBE BREEDING HABITAT, SASKATCHEWAN By TABLE 2—Numbers of Horned Grebe nests in relation to pond size and permanency type (1974-1975) Numbers of nests! and ponds! in each permanency type Baral Type | Type 3 Type 4 Type 5 Nests size per (ha) nests ponds nests ponds nests ponds _ nests ponds pond? < 0.21 0 196 0 326 7 227 3 41 0.04 0.21—-0.40 0 13 0 64 8 118 21 81 0.15 0.41-0.81 0 2 0 29 16 114 30 94 0.22 0.82-1.62 0 0 0 8 8 55 31 96 0.26 1.63-3.24 0 0 0 1 2 11 16 49 0.30 > 3.24 0 0 0 0 1 4 i, 18 0.36 Totals 0 211 0 428 42 529 108 379 0.17 'Figures for 1974 and 1975 pooled. 2Average for pooled Types 4 and 5 ponds. brood trap for grebes that nested on them. In 1974, 13% of nests occurred on Type 4 ponds and 87% on Type 5 ponds. Figures for 1975 were 35% and 65% for Types 4 and 5 ponds, respectively. In both years there was unexpected high use (P< 0.001) of Type 5 ponds based on pro- portions of those present. As with birds observed during counts, the distribution of nests among the 12 sections was correlated with numbers of Types 4 and 5 ponds (r = 0.8; P< 0.002). Pond occupancy rate by nesting Horned Grebes increased with pond size (Table 2). It was comparatively low for ponds up to 0.2 ha and increased gradually for larger ponds. The relationship was strongest for Type 4 ponds. The probability of a Type 4 pond having patches of open water increased with size, and I believe this influenced selection by grebes. All Type 5 ponds had some open water; size would be less important in these. One pond had more than one nest. This was a 7.8-ha, Type 5 pond with two nests. Munro’s (1941) observations in British Columbia sug- gested that two or more nesting pairs on one pond were more common there. He observed three pairs on a 2-ha pond in two successive years and five nests one year ona 30-ha lake. In western Saskatchewan, Renaud and Renaud (1975) reported four breeding pairs on a 8.1-ha pond. Stewart (1975) stated that Horned Grebes in North Dakota usually nested as solitary pairs on the smaller ponds or as widely scattered pairs on the larger lakes. An exception was five nests on a 17.4-ha pond. Fjeldsa (1973b) found that coexistence (two or more pairs) seldom occurred on “unfertile” ponds under 5 ha in Iceland. In contrast, it was relatively common on “fertile” ponds over | ha. He showed that, on the average, grebe pairs had exclusive use of areas many times larger than that needed for successful breeding, and suggested that territorial behavior brought about an overdispersal of pairs. This also seemed to be the case on my area and the comparatively high density of suitable ponds allowed pairs to nest solitarily. The presence or absence of tree growth around ponds had no apparent effect on the choice of ponds by nesting Horned Grebes. Ratios of treed ponds selected to those present did not differ significantly in 1974 (P > 0.05) or in 1975 (P> 0.5). From data based on nest distribution, nesting Horned Grebes did not show a preference for pasture ponds as did birds in the pre-nesting population either in 1974 (P >0.8) or in 1975 (P> 0.1). The tendency for nesting pairs to disperse may be stronger than their attraction to ponds in uncultivated areas. Nest Sites The sites of 92 nests were recorded in 1975; 40% of the nests were supported by flooded willows (Salix spp.) and 10% by dead cattail (Typha latifolia). The others, 50%, were floating or loosely anchored structures in open sites, sometimes surrounded by new growth of whitetop (Scolochloa festucacea) or, less com- monly, sedge (Carex atherodes) later in the season. Water depth at 50 nests in 1975 averaged 86cm (range, 25 to 140cm). This is a 376 conservative estimate because a few were not recorded as they were too deep to reach by wading and were not visited with a canoe. Depth at 11 nests measured by Stewart (1975) in North Dakota averaged 41 cm (range, 15 to 122 cm). The comparatively high average depth for my sample probably reflects the relatively high water levels in the ponds that year. Conclusions Throughout the breeding season, Horned Grebes used the more permanent ponds almost exclusively. Ponds over 0.2 ha were also favored, probably because they contained more open water. No other habitat factor appeared to affect the distribution of nesting grebes. Land use practices such as drainage and filling tend to affect adversely temporary wetlands ata higher rate than permanent wetlands (Kiel et al. 1972). Thus, compared with many bird species associated with wetlands, the Horned Grebe enjoys a relatively secure habitat. Although a few small temporary ponds on the study block had been destroyed, there was no evidence that any of the permanent ponds had been altered significantly. This small area, however, cannot be considered representative of the Canadian parklands. Elsewhere, wetlands, including per- manent ponds, are being destroyed at a higher rate (Kiel et al. 1972). Little is known about the effects of agri- cultural chemicals on these pothole ecosystems. These also need investigation because, in the long term, they may be just as damaging as physical alteration. Acknowledgments I thank J. H. Patterson, J. Fyeldsa, J. B. Gollop and the referees for their helpful reviews of the manuscript. Literature Cited Bent, A. C. 1963. Life histories of North American diving birds. Dover, New York. Bird, R. D. 1961. Ecology of the aspen parkland of western Canada. Canada Department of Agriculture Publication 1066. THE CANADIAN FIELD-NATURALIST Vol. 91 Driver, E. A. 1977. Chironomid communities in small prairie ponds: some characteristics and controls. Fresh- water Biology 7: 121-133. Driver, E. A. and D.G. Peden. 1977. The chemistry of surface water in prairie ponds. Hydrobiologia 53: 33-48. DuBois, A.D. 1919. An experience with horned grebes (Colymbus auritus). Auk 36: 170-180. Dwyer, T. J. 1970. Waterfowl breeding habitat in agri- cultural and nonagricultural land in Manitoba. Journal of Wildlife Management 34: 130-136. Evans, C.D. and K.E. Black. 1956. Duck production studies on the prairie potholes of South Dakota. United States Fish and Wildlife Service Special Scientific Report, Wildlife Number 32. Faaborg, J. 1976. Habitat selection and territorial behavior of the small grebes of North Dakota. Wilson Bulletin 88: 390-399. Fjeldsa, J. 1973a. Feeding and habitat selection of the horned grebe, Podiceps auritus (Aves), in the breeding season. Videnskabelige Meddelelser fra Dansk Natur- historisk Forening 1 Kjobenhavn 136: 57-95. Fjeldsa, J. 1973b. Territory and the regulation of popula- tion density and recruitment in the horned grebe Podiceps auritus arcticus Boje, 1822. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 1 Kjoben- havn 136: 117-189. Kiel, W. H., Jr., A.S. Hawkins, and N. G. Perret. 1972. Waterfowl habitat trends in the aspen parkland of Manitoba. Canadian Wildlife Service Report Series Number 18. Martin, A. C., N. Hotchkiss, F. M. Uhler, and W. S. Bourn. 1953. Classification of wetlands of the United States. United States Fish and Wildlife Service Special Sci- entific Report, Wildlife Number 20. Millar, J. B. 1976. Wetland classification in western Cana- da. Canadian Wildlife Service Report Series Number 37. Mitchell, J.. H.C. Moss, and J.S. Clayton. 1947. Soil survey of southern Saskatchewan from Township | to 48 inclusive. University of Saskatchewan Soil Survey Report 12. Munro, J. A. 1941. Studies of waterfowl in British Colum- bia. The grebes. Occasional Papers of the British Columbia Provincial Museum Number 3. Renaud, W.E. and D.H. Renaud. 1975. Birds of the Rosetown-Biggar District, Saskatchewan. Saskatchewan Natural History Society Special Publication 9. Smith, A. G. 1971. Ecological factors affecting waterfowl production in the Alberta parklands. United States Fish and Wildlife Service Resource Publication 98. Stewart, R.E. 1975. Breeding birds of North Dakota. Tri-College Center for Environmental Studies, Fargo, North Dakota. Received 3 May 1977 Accepted 6 September 1977 The Structure and Rate of Growth of the Rhizomes of Some Forest Herbs and Dwarf Shrubs of the New Brunswick - Nova Scotia Border Region DOUGLAS G. SOBEY! and PETER BARKHOUSE? Department of Biology, Mount Allison University, Sackville, New Brunswick EOA 3C0 1Present address: Stranmillis College, Belfast BT9 SDY, Northern Ireland 2Present address: Canadian Wildlife Service, Sackville, New Brunswick EOA 3C0 Sobey, Douglas G. and Peter Barkhouse. 1977. The structure and rate of growth of the rhizomes of some forest herbs and dwarf shrubs of the New Brunswick — Nova Scotia border region. Canadian Field-Naturalist 91(4): 377-383. Rhizome growth was examined in 41 herbs and dwarf shrubs of hardwood and mixed forests in the New Brunswick — Nova Scotia border region of Canada. For each species the morphology of rhizome growth was examined and the rate of growth (cm/yr) was estimated. Thirty-four of the species had either surface or underground rhizomes and in 32 of these ramification or division of plants was observed. Two of the species that lacked rhizomes were able to spread by means of horizontal roots on which adventitious buds developed. The maximum rates of rhizome or root growth varied from almost nil for some species to rates approaching 1 m/yr for others. The frequency of branching also varied although this was not precisely determined for many species. Examination of the results indicates that most of the species of slower rhizome growth are characteristic of the climatic climax fern-herb forests of the region, whereas many of the species of faster rhizome growth are more characteristic of the successional stages after fire, logging, or other disturbances. The adaptive significance of these findings is discussed. Key Words: rhizome structure, rhizome growth, vegetative spread, forest herbs, maritime provinces. Vegetative reproduction by means of the growth and branching of horizontal rhizomes has long been recognized as an important method of plant propagation, but comparatively little is known about such reproduction in natural habitats. Several European workers have investigated different aspects of vegetative propagation in meadow and forest species (Oinonen 1967a, b, 1968, 1971; Harberd 1961, 1963; Tamm 1972a, b). North American studies include those of Holm (1925), Whitford (1949, 1951), Struik (1965) and Bell (1976). Some of the above studies included estimates of the rates of rhizome growth of particular species but none attempted to examine and compare the rates in a large number of species from the same type of habitat. This was the purpose of the work reported here which was carried out near Sackville, New Brunswick in the summer of 1969. It was intended to examine rhizome growth among the plants comprising the ground flora of local forests. Forty-one of the 53 species considered eligible were examined and for each of these an attempt was made to determine the morphology of rhizome growth and the rate of such growth in centimetres per year, including where possible the extent of branching. Since the study was intended as a broad survey in a field that has been largely unexplored, there are gaps in the data and most of the species would benefit from more detailed separate investigations. It should also be noted that onlya few specimens of each plant species were examined, and though they are considered typical of the habitat in which they are found, they are not necessarily representative of the species throughout its range or in all types of habitat. Methods Thirty-two of the species were examined ina forest along the Upper Walker Road, about 4.5 km from Sackville, Westmorland County, New Brunswick. The site contained extensive stretches of young hardwood trees and has a long history of selective logging. Sugar maple (Acer saccharum), beech (Fagus grandifolia), and yellow birch (Betula alleghaniensis) were the principal tree species with occasional red maple (Acer rubrum), paper birch (Betula papyrifera), red spruce (Picea rubens), and balsam fir (A bies balsamea), the latter two occurring as dominants in some areas. The soil was a podzol with a loose well-humified litter layer. Eight species (Polystichum acrostichoides, SHH 378 Athyrium filix-femina, Streptopus roseus, Poly- gonatum pubescens, Medeola virginiana, Actaea pachypoda, Mitchella repens, and Prenanthes altissima) were collected from a mature hard- wood forest near Fenwick, Cumberland County, Nova Scotia. At this site occasional hemlock (Tsuga canadensis) occurred, along with the dominants sugar maple, beech, and yellow birch. The soil at this site was also a podzol. One species, Pyrola elliptica, was collected along a roadside near Sackville. Each species was first examined in situ. Speci- mens were then dug up with the underground parts of some species being mapped. Further measurement and examination of the specimens took place in the laboratory. Though the rate of rhizome growth was determined accurately for some species, for others it was only estimated and for some it could not be determined during the time of the study. The methods used to assess the rate of growth of horizontal rhizomes or stolons were the following: (1) Species with persistent (i.e., wintergreen) leaves along their rhizomes frequently had features which distinguished one year’s increment from another. All six clubmosses had clusters of either shorter leaves or scales on their aerial shoots and/or on their horizontal above-ground rhizomes marking the division between two year’s growth (Figure 1). The amount of growth of Lycopodium lucidulum, L. clavatum, L. annotinum, L. obscurum, and L. flabelliforme was thus measured exactly for several years back, and that of L. trista- chyum was estimated. In L. /ucidulum the persistent sporangial cases of recent years corroborated the above leaf evidence. In other species with persistent leaves along hori- zontal stems (Linnaea borealis, Mitchella repens, Gaultheria hispidula) slight differences in the color and size of leaves and the color and shape of stems were used to distinguish the current year’s growth from that of previous years (Figure 1). Thus a fairly accurate measure of the amount of new growth and sometimes that of preceding years was obtained. (2) The remains of old shoot or leaf bases along a rhizome or of scars marking the position of old shoots were used to estimate the rate of growth of many plants including the ferns, the stipe bases of which persist for many years (Figure 1). By counting the number of dead stipe bases along a rhizome and counting the number of fronds produced per year at shoot points on living plants, the rate of horizontal growth was estimated for the following species: THE CANADIAN FIELD-NATURALIST Vol. 91 Osmunda claytoniana, Polystichum acrostichoides, Dryopteris spinulosa, D. phegopteris, Athyrium filix- femina, and Pteretis pensylvanica. The presence of scars or shoot remnants were also used to estimate the growth rate of Smilacina racemosa (Figure 1), Streptopus roseus, Polygonatum pubescens, Clin- tonia borealis, Aster acuminatus (Figure 1), Ana- Phalis margaritacea, Trillium erectum (Figure 1), Trillium undulatum (Figure 1), and Galium triflorum. Sometimes shoot points continue to produce a shoot for several years while new shoot points are continually added at the anterior growing end of a rhizome. The distance between such shoots was assumed to equal the annual increment and, along with supporting information (see below), was used to estimate the growth rates of Aralia nudicaulis and Maianthemum canadense. (3) In Medeola virginiana and Trientalis borealis (Figure 1), the position of the following year’s shoot was marked by a tuber which developed at the end ofa rhizome in late summer. The distance between the current year’s shoot and this tuber represents the annual rhizome growth. Determined in somewhat the same way was Pteridium aquilinum in which future year’s fronds were present in various stages of develop- ment, the evidence suggesting that the distance between the incipient fronds represents an annual increment. (4) For some species (Cornus canadensis, Gaultheria procumbens, Coptis trifolia, Oxalis montana, and Maianthemum canadense) the rhizome produced in the current year was often lighter in color than that produced during previous years, and this was used to estimate the growth rate where there was no other evidence. (5) Internodal lengths on the horizontal rhizome (i.e., the distance between scales or leaves) aided with Aralia nudicaulis and Maianthemum canadense. Shorter internodes were presumed to mark the cooler days of fall and thus the point of division between two years’ growth. No estimate was obtained for the rate of growth of Pyrola elliptica, Epilobium angustifolium and Aralia hispida, the last two of which spread vegetatively by the horizontal growth of roots (confirmed by micro- scopic examination of cross-sections). Results Morphology of Rhizome Growth Of the 41 species examined, 34 had either surface or underground rhizomes (the seven not possessing rhizomes were Prenanthes trifolio- lata, P. altissima, Actaea pachypoda, Mono- tropa uniflora, Cypripedium acaule and the two species spreading by the horizontal growth of 1977 SOBEY AND BARKHOUSE: RHIZOMES, MARITIMES 379 Trillium undulatum CURRENT YEARS ne A Ain | is] PREVIOUS YEAR'S GROWTH iN / Lycopodium annotinum aN Trillium SPORANG/A 29) erectum ee! Lycopodium lucidulum S : milacina racemosa J 010 SHOOT SCARS Gaultheria hispidula Aster acuminatus CURRENT PoInT of SHOOT E PREVIOUS YEAR’S SHOOT SHOOT PoINT of 2 YEARS AGO GROWING RHIZOME TIP Aralia hispida Dryopteris spinulosa DEAD STIPE Trientalis borealis POINT of NEXT CURRENT YEAR'S SHOOT SHOOT THICK RHIZOME THIN RHIZOME in some of the species examined. All are rhizomes or surface stems are roots. For Lycopodium annotinum a cluster of short scale- h is shown. Bar scale for each figure is | cm. FiGuRE 1. Structures associated with vegetative spread except the underground parts of Aralia hispida which leaves marking the division between two years’ growt 380 roots — Epilobium angustifolium and Aralia hispida). In 32 of these 34, vegetative spread (1.e., ramification or division of plants) was observed (Table 1); the two not showing clear-cut rami- fication were Trillium erectum and T. undu- latum. In those species spreading by means of the growth and branching of a horizontal rhizome, the older posterior parts of the rhizome grad- ually die and decay, leading to the fragmen- tation of the original plant. In some species (Lycopodium lucidulum, L. annotinum, L. clavatum, Gaultheria hispidula, Galium _ tri- florum, Mitchella repens, Linnaea borealis) the rhizome (or stolon) initially developed as a prostrate stem on the soil surface which usually became covered with surface litter after the first year. In the remaining species the rhizome developed below the surface, generally in the upper few centimetres of the humus layer, though in Pteridium aquilinum it was deep (about 15 cm) in the eluviated horizon of the mineral soil. The shape and form of the rhizome and the morphology of branching showed some varia- tion but a detailed examination of this aspect was beyond the scope of this study. In general the rhizomes were of two types: some were long and slender with the nodes (marked by fine scales or leaves) at least several centimetres apart (e.g., Figure |, Aster acuminatus); others were short and stout, with the nodes, and frequently the points of origin of shoots, bunched together (e.g., Figure 1, Smilacina racemosa). In three species (Dryopteris spinulosa (Figure 1), Pteretis pensylvanica, Streptopus roseus), both slender and stout rhizomes were noted. In two of the species (Medeola virginiana and Trientalis borealis) a special form of rhizome growth was noted (Figure 1). Both slender rhizomes and aerial shoots develop in the early part of the growing season from a tuber. In late summer horizontal growth of the rhizome(s) ceases and a new tuber develops at the tip of each rhizome. The old tuber and rhizome decay over winter and the following season’s shoot develops from the new tuber. Though the Trillium species were not observed to spread vegetatively, they did have under- ground rhizomes about 10cm deep in the mineral soil (Figure 1). These had very short THE CANADIAN FIELD-NATURALIST Vol. 91 internodes and were tuber-like. Tiny tuber-like structures without shoots attached were found adjacent to larger shoot-producing T. erectum rhizomes. They appeared to have originated from the larger rhizome though no connection was observed. Rate of Horizontal Growth of Rhizomes, Stolons, and Roots The rate of rhizome growth varied from almost nil for some species to rates approaching 1m per year for others (Table 1). There was variation among plants of the same species and also from year to year for the same plant. A plotting of the estimated maximum rates of rhizome growth (Figure 2) permits comparison between species. The frequency of branching of the horizontal rhizomes or roots also showed wide variation (Table 1), but for many species it could not be precisely determined from a single season’s work. Discussion The rates of rhizome growth in Table | are generally similar to values obtained for the same species in other studies. Whitford (1951) fe- corded a growth rate of 0.73 in/yr (1.9 cm) for Athyrium filix-femina rhizomes (slightly in excess of that recorded in this study), and 0.94 in/yr (2.4cm) for Smilacina racemosa (which falls within the range of 1-3cm in Table 1). Anderson and Loucks (1973) in a detailed study of Trientalis borealis examined 93 plants and recorded rhizome lengths ranging from 5 cm to greater than | m withan average of about 30 cm; in this study lengths of up to 80 cm were recorded. Oinonen (1967a) estimated an annual radial increase of 17.9 cm for Pteridium aquilinum clones in Finland, which falls within the 15- to 30-cm range observed in this study, and 15.1 cm/yr for Lycopodium complanatum (Oinonen 1967b); the values for L. flabelliforme (which Fernald (1950) considers to be a variety of L. complanatum) were 17-31 cm. He also re- corded (Oinonen 1968) a range of 3 to 86 cm in the annual growth of shoots of L. clavatum (8-74 cm were observed in this study), and 2-58 cm for L. annotinum (5-45 cm were re- corded in this study). In Massachusetts Primack (1973) recorded the following ranges in the rhizome growth of five Lycopodium species: L. 1977 TABLE |—Data on vegetative spread for the 41 forest herbs and dwarf shrubs examined SOBEY AND BARKHOUSE: RHIZOMES, MARITIMES 381 Species Lycopodium lucidulum L. annotinum L. clavatum L. obscurum L. flabelliforme L. tristachyum Osmunda claytoniana Pteretis pensylvanica® Dryopteris phegopteris D. spinulosa* Polystichum acrostichoides Athyrium filix-femina Pteridium aquilinum Clintonia borealis Smilacina racemosa? Maianthemum canadense Streptopus roseus® Polygonatum pubescens” Medeola virginiana? Trillium erectum® T. undulatum Cypripedium acaule Coptis trifolia Actaea pachypoda Oxalis montana Epilobium angustifolium Aralia hispida A. nudicaulis Cornus canadensis Monotropa uniflora Pyrola elliptica Gaultheria procumbens G. hispidula Trientalis borealis Galium triflorum Mitchella repens Linnaea borealis Aster acuminatus Anaphalis margaritacea Prenanthes trifoliolata P. altissima Vegetative spread (i.e. ramification observed) + +++ + +++ ¢ + 4+ 4+ ¢ + + ~~ None observed None observed + None observed + + + None observed None observed None observed Observed range in growth rate or max. growth rate of horizontal rhizomes, roots or stolons (cm/yr) 1-3 5-45 8-74 13-20 17-31 ca. 15 (max.) < 4 (max.) <1 (max.) 14-3 <1; <4 (max.) <1 (max.) <1 (max.) 15-30 6-10 1-3 ca. 15-30 <1; 44 (max.) 1-3 2-8 0.4-0.6 0.4-0.5 Nil 5-(15?) Nil 10 (max.) Not determined Not determined 25-80 >18 (max.)? Nil Not determined >10 (max.)? 2-7 80 (max.) <5 (max.) <10 (max.) 11-48 1-25 2-20 Nil Nil Frequency of rhizome branching Irregular—|! branch/3-12 or more years Frequent (ca. 1/yr) 1 branch/3-4 yr 1 branch/yr Irregular but frequent (ca. 1 branch/ yr) Not determined 1 branch/75 yrs Not determined 1 branch/6-11 yrs Not determined Not determined (but infrequent) Not determined (but infrequen<’ 1 branch/yr 1 branch/4-10 yrs None observed Not determined (appears frequent 1 branch/ 1-3 yr?) Not determined None observed None observed None observed None observed None observed Not determined (appears frequent 1 branch/ yr?) None observed Not determined Not determined but appears frequent Not determined Potentially 1 branch/yr but much less—1 branch/4-5 yr(?) Appears frequent None observed Not determined Not determined, irregular Not determined, but frequent Not determined Not determined Irregular but frequent (1 branch/ yr?) (1 branch/ 3-4 yr?) Not determined Not determined but frequent None observed None observed *Pteretis pensylvanica, Streptopus roseus, and Dryopteris spinulosa (see Figure |) had both thick and thin rhizomes. For Pteretis the growth rate of only the thick thizome was determined; for Dryopteris and Streptopus both rates were determined and that of the thick is listed first. "Actual ramification was not observed in Polygonatum pubescens, Medeola virginiana, and Smilacina racemosa but the presence of lateral buds on the rhizomes suggest a capacity for such ramification. “Special tuber-like structures were observed near Trillium erectum rhizomes (see main text). 382 centimetres re) 5 10 20 30 40 Lycopodium Maianthemum Aster acuminatus Anaphalis margaritacea Lycopodium obscurum Cornus canadensis Coptis trifolia Lycopodium tristachyum Gaultheria procumbens Clintonia borealis Oxalis montana Mitchella repens Medeola virginiana Gaultheria hispidula Galium triflorum Streptopus roseus Dryopteris spinulosa Lycopodium lucidulum Smilacina racemosa Polygonatum pubescens Dryopteris phegopteris Athyrium filix-temina Pteretis pensylvanica Polystichum acrostichoides Osmunda-_claytoniana Trillium erectum Trillium undulatum THE CANADIAN FIELD-NATURALIST Vol. 91 Trientalis borealis Aralia nudicaulis Lycopodium clavatum Linnaea borealis Lycopodium annotinum flabelliforme Pteridium aquilinum canadense FIGURE 2. The estimated or observed maximum annual rates of horizontal rhizome growth for 33 of the 41 species examined. Of the remaining eight species, horizontal growth was not observed in five (Cypripedium acaule, Monotropa uniflora, Actaea pachypoda, Prenanthes altissima, and P. trifoliolata) and no estimate was obtained for Pyrola elliptica and the two species spreading by horizontal roots (Epilobium angustifolium and Aralia hispida) both of which appear to spread rapidly. flabelliforme (26-50 cm/yr), L. clavatum (48- 103), L. annotinum (22-45), L. obscurum (13- 21), L. lucidulum (1.5-2.2) — all of which, to varying degrees, overlap the ranges recorded in Table 1. The wide range observed among the species in the rates of rhizome growth and branching is not surprising in view of the taxonomically diverse group of plants examined. Among particular taxa there is some consistency: the clubmosses, with one exception, are among the species with faster growing rhizomes (all have maximum rates of rhizome growth greater than 15 cm/yr). The exception, L. lucidulum, has a maximum rate of only 3 cm. In contrast the ferns (like the clubmosses, spore-producing plants) with the exception of Pteridium aquilinum, are plants of slow rhizome growth, mostly under | cm/yr. Pteridium by contrast has a maximum rate of 30 cm. It is more informative to group the plants according to the community in which they occur rather than according to taxonomics. Many of the plants of slower or no rhizome growth (Lycopodium lucidulum, Dryopteris phegop- teris, D. spinulosa, Polystichum acrostichoides, Athyrium filix-femina, Smilacina racemosa, Streptopus roseus, Medeola virginiana, Clin- tonia borealis, Oxalis montana, Monotropa uniflora, Mitchella repens, Trillium undulatum, T. erectum) are characteristic of the sugar maple- beech-hemlock fern-herb forest which is considered to be the climatic climax forest on the better soils in New Brunswick — Nova Scotia. In such a supposedly stable climax habitat plants theoretically have an unlimited time in which to colonize the habitat and this may be the reason why many species have not evolved a more rapid rate of rhizome spread. By contrast, many of the species of more rapid rhizome growth (Lycopodium annotinum, L. clavatum, L. obscurum, L. flabelliforme, L. tristachyum, Cornus canadensis, Linnaea bore- alis, Pteridium aquilinum, Anaphalis margari- tacea, and Aster acuminatus (plus Epilobium angustifolium and Aralia hispida whose rates of spread were undetermined but would appear to be rapid) are species more characteristic of the 1977 SOBEY AND BARKHOUSE: early to late successional stages after fire, logging, or other disturbances. In such succes- sional communities where environmental condi- tions are slowly but continually changing there is a limit to the time that a seral species will find favorable conditions enabling it to occupy a site. Under such circumstances rapid rhizome growth and spread may have evolved as an adaptive advantage permitting extensive colonization before conditions became unfavorable. More precise estimates may even reveal a gradation in the rate of rhizome growth among the early to late successional herbs. The taxonomic anomaly mentioned above is now seen to be explained by the ecological status of the species. The only slow-growing clubmoss, L. /ucidulum, is also the only clubmoss characteristic of the climax forest; Pteridium, the fast-spreading fern, is the only fern examined characteristic of the early succes- sional stages. This study has been a rather broad survey of rhizome growth among a group of plants from the same habitat. Our understanding of most of the species might benefit from more detailed examination. Also other species and other habitats deserve investigation. It may well be found that the difference in the rates of rhizome growth between species of seral and climax communities suggested by this study is a feature not only of forests, but also of other com- munities. Acknowledgments We are indebted to Hinrich Harries of Mount Allison University for suggesting the topic for research, and for guidance and help during the study. We thank him and also David Smith and David Park of the Botany Department of the Queen’s University of Belfast, for helpfully reading and criticizing the paper. Literature Cited Anderson, R. C. and O. L. Loucks. 1973. Aspects of the biology of Trientalis borealis Raf. Ecology 54: 798-808. RHIZOMES, MARITIMES 383 Bell, A. 1976. Computerized vegetative mobility in rhizo- matous plants. Automata, Languages, Development (Edited by A. Linden-Mayer and G. Rozenburg). North- Holland Publishing Co., Amsterdam. p. 3-14. Fernald, M. L. 1950. Gray’s manual of botany. 8th edition. American Book Company, New York. Harberd, D. J. 1961. Observations on population structure and longevity of Festuca rubra L. New Phytologist 60: 184-206. Harberd, D. J. 1963. Observations on natural clones of Trifolium repens L. New Phytologist 62: 198-204. Holm, T. 1925. Hibernation and rejuvenation, exemplified by North American herbs. American Midland Naturalist 9: 439-512. Oinonen, E. 1967a. Sporal regeneration of bracken (Preri- dium aquilinum (L.) Kuhn.) in Finland in the light of the dimensions and the age of its clones. Acta Forestalia Fennica 83.1: 1-96. Oinonen, E. 1967b. Sporal regeneration of ground pine (Lycopodium complanatum L.) in southern Finland in the light of the size and the age of its clones. Acta Forestalia Fennica 83.3: 1-75. (In Finnish, English summary.) Oinonen, E. 1968. The size of Lycopodium clavatum L. and L. annotinum L. stands as compared to that of L. complanatum L. and Pteridium aquilinum (L.) Kuhn stands, the age of the tree stand and the dates of fire on the site. Acta Forestalia Fennica 87: 1-53. (In Finnish, English summary.) Oinonen, E. 1971. The time table of vegetative spreading in oak fern (Carpogymnia dryopteris (L.) Love & Love) and May-lily (Maianthemum bifolium (L.) F. W. Schmidt) in southern Finland. Acta Forestalia Fennica 118: 4-37. Primack, R. B. 1973. Growth patterns of five species of Lycopodium. American Fern Journal 63: 3-7. Struik, G. J. 1965. Growth patterns of some native annual and perennial herbs in southern Wisconsin. Ecology 96: 401-420. Tamm, C. O. 1972a. Survival and flowering of some peren- nial herbs. II. The behaviour of some orchids on permanent plots. Oikos 23: 23-28. Tamm, C. O. 1972b. Survival and flowering of perennial herbs. III. The behaviour of Primula veris on permanent plots. Oikos 23: 159-166. Whitford, P. B. 1949. Distribution of woodland plants in relation to succession and clonal growth. Ecology 30: 199-208. Whitford, P. B. 1951. Estimation of the age of forest stands in the prairie-forest border region. Ecology 32: 143-147. Received 13 April 1977 Accepted 13 September 1977 New Vascular Plant Records on Pelee and East Sister Islands, Essex County, Ontario CRAIG A. CAMPBELL! and A. A. REZNICEK2 1421 King Street North, Waterloo, Ontario N2J 3Z4 2Department of Botany, Erindale College, University of Toronto, Mississauga, Ontario LSL 1C6 Campbell, C. A. and A. A. Reznicek. 1977. New vascular plant records on Pelee and East Sister Islands, Essex County, Ontario. Canadian Field-Naturalist 92(1): 384-390. Pelee Island, the southernmost part of Canada, has a diverse and very interesting flora containing many species rare or absent elsewhere in Canada. Reported as new to Canada are seven species of vascular plants: Carex davisii, Carex divulsa subsp. leersti, Sedum telephioides, Euphorbia obtusata, Myosotis macrosperma, Lycopus virginicus var. virginicus, and Senecio glabellus. These are briefly discussed. In addition, we report on the status of 13 very rare species known previously from the area: Camassia scilloides, Spiranthes magnicamporum, Celtis tenuifolia, Chenopodium foggii, Thalictrum dasycarpum, Corydalis flavula, Cydonia oblonga, Ammannia coccinea, Chaerophyllum procumbens, Phacelia purshii, Conobea multifida, Eupatorium altissimum, and Eclipta prostrata. Fifty-five other species that are additions to the published flora of the Erie Archipelago, Pelee Island, and East Sister Island are listed as well. Key Words: vascular plants, floristics, Erie Archipelago, Ontario, Canada, endangered species, phytogeography. Field work on Pelee Island, Essex County, Ontario, during the past 10 years has resulted ina number of discoveries of considerable floristic interest. Several species not previously reported for Ontario or even for Canada have been recorded, although the Erie Islands have been quite extensively botanized (Macoun 1893; Core 1948; Duncan 1973). The first botanist to collect extensively on Pelee Island was John Macoun, Canadian Dominion Botanist, in 1882 (Macoun 1893). Later came C. K. Dodge, who produced the first list of Pelee Island plants (1914), and then Core (1948). The collections of Wilfred Botham, of Cottam, Ontario were largely the basis for Boivin’s (1953) additions to Core’s list. Recent work by Stuckey (1968), Duncan and Stuckey (1970), and Duncan (1973), as well as by Stuckey and Duncan, currently updating the work of Core, has added considerably to the published flora. Our paper reports species in addition to those recorded in these lists. Pelee Island in particular has a wide diversity of plant habitats and a large flora for sucha small area. The island is situated in the western end of Lake Erie, in extreme southern Canada between 41°43’ to 41°50’N, and 82°37’ to 82°42’W. It is the largest of the Erie archipelago islands: 4083 h (10085 acres). East Sister Island, also part of Pelee Township, is approximately 13 km (8 mi) west of Pelee Island, at 41°49’ N, 82°50’ W; it is some 26 h (65 acres) in size, of which only about 14.6 (36) have been above water recently. The warm climate of Pelee Island is largely a result of the moderating effects of Lake Erie; the mean annual temperature is 9.6°C (49.3°R), several degrees higher than that of the mainland of Essex-Kent Counties. The average growing season (free of killing frosts) is 192 days. Droughty conditions do occur as a result of lighter rainfall than on the mainland (Core 1948). Pelee Island has relatively low rainfall, slightly under 75 cm (30 in) per year. The flora of Pelee Island has strong affinities with the Ohio-Mississippi River lowlands, and this affinity is reflected by some of the most abundant trees on the island, e.g., Celtis occi- dentalis, Acer nigrum, Fraxinus quadrangulata, Quercus macrocarpa, and Populus deltoides. The north shore of Ohio was mapped as having small prairie outliers by Transeau (1935); prairie species are fairly well represented on Pelee Island. Also, owing to its floristic affinities with the mid-west, Pelee Island, and especially the southern points of it (Figure 1) are much more similar to Point Pelee than to the Niagara Peninsula, despite the limestone bedrock. Few elements of the eastern Carolinian and/or Appalachian flora occur on Pelee Island; the Tulip-tree (Liriodendron tulipifera) and Mag- nolia acuminata, for example, are missing from the Erie Islands. The flora of these islands is of continual interest because of the southern species which seem to shift northward and thus first reach Canada on this archipelago. Because of the warm climate of Pelee Island, 384 ID PELEE ISLAND Point —oualy FIGURE 1. Map of Pelee Island showing major collecting locations. several cultivated species which persist and slowly spread on Pelee Island do so rarely or not elsewhere in Canada. These have not been discussed in the list. They include Rosa multi- flora, Lonicera japonica, and Prunus persica. The flora of Pelee Island is also somewhat paralleled by the fauna. The Small-mouthed Salamander (Ambystoma texanum), the Blue Racer (Coluber constrictor foxi), the Red-bellied Woodpecker (Melanerpes carolinus), and Blan- chard’s Cricket Frog (Acris crepitans blan- chardi) occur in Canada as well established populations on Pelee Island only. These species are widespread in the central Mississippi-Ohio lowlands, but very rare or absent elsewhere north of the international boundary. Our floristic additions are recorded in the form of an annotated list, dealing first with vascular plants new to Canada, and second with recent observations of rare species. A short dis- cussion of the floristic and phytogeographic significance of each species is provided. A table summarizing other additions to the flora of Pelee CAMPBELL AND REZNICEK: VASCULAR PLANT RECORDS, S.W. ONTARIO 385 and East Sister Islands and including whether the species are new to the flora of the Erie Archipelago is also included. Our major collecting sites are shown in Figure 1; others are specified in the annotated list which follows. Habitats are briefly noted; more details in connection with the present paper can be obtained from the Canadian Botanical Association’s Survey of the Rare and En- dangered Plants of Canada. Annotated List of Species New to Canada Carex davisii Schwein. & Torr. Essex County, 9.6 km (6 mi) northeast of Amherst- burg along Canard River, occasional at edge of deciduous woods, 24 June 1955, Calder and van Rens 15928 (DAO,TRT) (sub C. formosa). Pelee Island, Mill Point, south side of main E-W road, edge of woods and cultivated field, 28 June 1975, Campbell and Harris (TRTE, WLU). This southern and mid-western species of lowland woods and meadows was previously recorded from nearby South Bass and Kelley’s Islands, Ohio (Core 1948) and sparingly from extreme southern Michigan (Voss 1972). The report of this species from Quebec (Marie-Victorin 1935) has been shown to be an error by Rouleau (1964). Carex divulsa Stokes subsp. Jeersii (Aschers & Graebn.) W. Koch (C. polyphylla Kar. + Kir., C. leersiana Rauschert, C. leersii F. W. Schultz). Along South Road at junction with Stone Road just west of Mill Point, Pelee Island, 8 August 1974, Reznicek, Campbell and Donaldson (TRTE). A Eurasian sedge, C. divulsa subsp. /eersii prefers calcareous soil (Jermy and Tutin 1968), as is the case on Pelee Island. The taxonomic status of the Carex muricata aggregate, to which this species belongs, is not yet fully clear. We follow here the nomenclature proposed by David and Chater (1977). Further discussion of this difficult complex of species is provided by Jermy and Tutin (1968), David and Kelcey (1974), and David (1976). Complete synonomy is listed in David (1976). Sedum telephioides Michx. Pelee Island, Verbeek savanna, dry limestone outcrop, pasture, 4 September 1974, Campbell and Donaldson (WLU). This species was growing with Corydalis flavula in open rocky savanna. Since it was observed growing here close to houses, it may well be an escape from cultivation. It was not reported for the Erie Islands by Core (1948) and Boivin (1953). Its native range is to 386 the south of Lake Erie (Clausen 1975), and it has not been previously reported from Canada. Euphorbia obtusata Pursh Essex County, River Canard, 17 June 1972, W. Botham (in herb. W. Botham). Pelee Island, Verbeek’s, Harris Road (north side), open oak savanna (grazed), 9 June 1974, Campbell and Donaldson (CAN, WLU). This is the first substantiated report of this species for Canada. Soper’s (1949) listing of this species from southern Ontario was based on the report by Billings (1862) (J. H. Soper, personal communications 1976). Billings’ report was based upon a specimen of E. platyphylla (Dore 1961). It is recorded by Gleason and Cronquist (1963) from Pennsylvania to Indiana. Myosotis macrosperma Engelm. (M. virginica var. macrosperma (Engelm.) Fern.) Pelee Island, Fish Point, woodland, 7 June 1970, J. K. Morton(WAT). Pelee Island, Brown’s (Middle) Point, low woods, 7 June 1974, Campbell (CAN, WLU). This large, wide-leaved plant of rich, low woods 1s quite distinct from its nearest relative in the Canadian flora, M. verna, which usually occurs in sunnier, drier habitats, as on Pelee Island limestone plains. Myosotis macrosperma is known from Ohio (Fernald 1950). Lycopus virginicus L. var. virginicus Pelee Island, Fish Point, edge of muddy field, 7 August 1974, Campbell, Reznicek, Donaldson (WLU); McCormick Woods, low wet woods with red ash, 7 August 1974, Reznicek (WLU); and Fish Point, moist sandy soil of dune slack, 7 August 1974, Reznicek, Campbell, Donaldson (TRTE). Although reported for Canada in the past, previous records by many authors all appear to be based on misidentified material of L. uniflorus. These three collections appear to be the first authentic Canadian material to be reported. Henderson (1962) discussed the features separating these two species, and has mapped L. virginicus as occurring mainly south of the glaciated zone in eastern North America. Its occur- rence on Pelee Island appears disjunct and similar to the distribution of Triosteum angustifolium (Duncan 1973), which also occurs on the island but otherwise largely south of glaciated territory. It has been recorded, however, from Green Island, Ohio (Core 1948). Senecio glabellus Poir. Butterweed Pelee Island, south marina, westernmost pond west of Bonnett’s, moist sandy and rocky shore of Lake Erie, 8 August 1974, Campbell, Reznicek and Donaldson (CAN). This annual or biennial is reported for southwestern Ohio, south-central Indiana, and southern Illinois by THE CANADIAN FIELD-NATURALIST Vol. 91 Gleason and Cronquist (1963). Stuckey (1975) records it as rare and local on the American Erie Islands and, in particular, in Perrys Monument Marsh on South Bass Island, where he considered it adventive from farther south. Recent Records of Rare Plants Camassia scilloides (Raf.) Cory Wild Hyacinth Pelee Island, Brown’s (Middle) Point, low rich open deciduous woods, 10 May 1974, Donaldson and Campbell P74-1 (CAN); and Fish Point, wet hedge- row, 7 August 1974, Campbell, Reznicek and Donald- son (WLU). Although this lovely spring lily was known from wet meadows on White Island in the Detroit River near Amherstburg (11 June 1882, J. Macoun, CAN), it has been recently collected in Canada only on the Erie Islands including Hen, Middle Sister, and North Harbour (Core 1948) and Middle Island (9-10 May 1939, H. Senn 1128 and J. H. Soper, TRT). This plant is vulnerable to picking and grazing and should be protected, or its extinction in Canada is probable. Spiranthes magnicamporum Sheviak Plains’ Ladies Tresses Pelee Island, Stone Road savanna, east side of road, dry grassy road verge, 25 September 1976, Diebollt, Francis and Campbell (WLU). This mid-western species, recently described (Shev- iak 1973), is known now in Ontario from a number of sites in the southwest (Catling 1976). Until this report, no Spiranthes were known on the Erie Islands, although Luer (1975) maps S. magnicamporum from the Ohio shore of Lake Erie. The Stone Road colony consisted of about 40 individuals. Celtis tenuifolia Nutt. Dwarf Hackberry Pelee Island, Fish Point, Lake Erie, 16 July 1958, L. J. Stock 348 (CAN) (sub C. occidentalis); and Fish Point, west side, near base of point, sand dunes, edge of Lake Erie (woods edge), 8 August 1974, Reznicek, Donaldson and Campbell (CAN, TRTE). The Dwarf Hackberries of the Great Lakes region have been designated C. tenuifolia var. soperi Boivin. Boivin (1967) recorded Canadian material only for the Grand Bend area, Lake Huron, as did Wagner (1974). This plant is new for the Erie Islands flora (see Core 1948). Chenopodium foggii Wahl Pelee Island, Fish Point, lagoon edge, sandy woods, 4 September 1974, Campbell and Donaldson (WLU); Lake Henry, clay and limestone rubble dike, 18 September 1975, Campbell, Reznicek and Donaldson PI-75-F15 (WLU, TRTE); and Verbeek’s savanna, 19 September 1975, Campbell, Reznicek and Donaldson PI-F41 (WLU). This uncommon native southern species has also 1977 been reported from Leeds County, Ontario and Pontiac County, Quebec (Wahl 1954) and from Lambton County, Ontario (Gaiser and Moore 1966), and no doubt occurs sparingly elsewhere in southern Ontario. Thalictrum dasycarpum Fisch. & Lall. Meadow-rue Pelee Island, Brown’s (Middle) Point, gravelly ditch, woods edge, 8 June 1974, Campbell and Donaldson (CAN, WLU). Considered a northwestern species in Ontario (B. Boivin, personal communication), it is known spar- ingly from southern Ontario. Our collection repre- sents an addition to the Erie Islands flora, although Core (1948) doubtfully reported it from South Bass Island, Ohio. Corydalis flavula (Raf.) DC. Yellow Corydalis Pelee Island, Verbeek savanna, Harris Road, rocky oak glade, 12 May 1974, Campbell P74-26 and Donaldson (CAN, WLU). This delicate fumitory is presently known in Canada only from Point Pelee and Pelee Island. On Point Peiee it has been collected as recently as 1970 (17 May, Morton, CAN,TRT, herb. J. K. Morton). It was first collected on Pelee Island by Macoun (Burgess 1889), who also collected it at Amherstburg, Ontario, (J. Macoun, 10 June 1882, MT'MG); and it was reported at Point Abino on the north-eastern shore of Lake Erie (Burgess 1889). Cydonia oblonga Mill. Quince Pelee Island, South Road (just west of south end, Stone Road), hedgerow along bush, 8 August 1974, Reznicek, Campbell and Donaldson (CAN, TRTE, WLU). The only other Canadian collection outside of cultivation is from Elgin County, along a creek at Aylmer, 4 June 1953, Montgomery and Shumovich 606 (TRT). Our record is an addition for the Erie Islands. Purple Ammannia coccinea Rottb. Essex County, Hillman Marsh, on mud flat, 42°03’N, 82°30’W, 7 September 1974, W. Botham 1704 (CAN); Pelee Island, Lake Henry, Lighthouse Point, mudflats, 19 September 1975, Reznicek and Diebolt and Campbell PI-75-F30 (DAO, WLU). The only other Canadian collections are from Lake Osoyoos, British Columbia: J. W. Eastham, 27 September 1939 (DAO); Calder and Saville 11519, 6 August 1953 (DAO). This is the first report of the species from eastern Canada; the collection by Campbell et al. is the first from the Erie Islands, although Core (1950) reported it from the Portage River swamps on the Ohio main- land. CAMPBELL AND REZNICEK: VASCULAR PLANT RECORDS, S.W. ONTARIO 387 Chaerophyllum procumbens (L.) Crantz Spreading Chervil Pelee Island, Brown’s Woods, 4 May 1971, Simp- son, Campbell and Pratt (WLU); Brown’s (Middle) Point, low rich deciduous woods, 11 May 1974, Campbell P74-27 and Donaldson(CAN); Mill Point, 12 May 1974, Donaldson and Campbell P74-23 (WLU); Stone Road, far to the east of Stone Road, 1 June 1974, W. Botham 1609 (CAN); and Verbeek savanna, grassy wooded glade by south fence, 5 May 1976, Perrin and Campbell PI-76-11 (CAN, WLU). This species was formerly known from Canada on White Island near Amherstberg, Ontario (moist places, 11 June 1882, J. Macoun, TRT), and from Kelley’s Island in Ohio (Core 1948). It was collected from Pelee Island in 1914 (Point Pelee Island, 27 May 1914, N. Tripp (MICH)), but not from there again until our collection of 1971. As is the case in Wisconsin (Curtis 1959), on Pelee Island this plant, although very local, may occur in dense colonies of hundreds. It is interesting that collections from Pelee Island include both C. procumbens var. procumbens and the pubescent-fruited var. shortii. The variety is appar- ently more western (Deam 1940; Fernald 1950; Gleason 1952), and Pelee Island is within the area of overlap of the two taxa. Phacelia purshii Buckley Miami-mist Pelee Island, Stone Road, | June 1974, W. Botham 1614 (CAN); Pelee Island, 22 June 1975, W. Botham 1762 (TRT). Reported by Dodge (1914) for Middle Sister and Chicken Islands in the Canadian Erie archipelago, this constitutes the first record of this plant from Pelee Island. These stations on the Erie islands constitute the only native Canadian occurrences; collections from a clover field in the Central Experimental Farm (W. T. Macoun, 3 June 1898, DAO; July 1898, CAN) are surely chance introductions. Conobea multifida (Michx.) Bentham Pelee Island, Stone Road, 13 June 1974, W. Botham 1672 (CAN); J. Harris lane behind barn, Lighthouse Point, muddy depressions, 19 September 1974, Rez- nicek and Diebolt (WLU); and Lighthouse Point, Lake Henry, mudflats, 19 September 1974, Reznicek, Campbell and Diebolt (WLU). This obscure member of the Scrophulariaceae has been collected previously on Pelee Island a number of times (Soper 1962) but nowhere else in Canada. Our collections extend its occurrence from the south parts of the island to the north end. It also occurs on Kelley’s Island, Ohio (Core 1948). Eupatorium altissimum L. Yall Boneset Essex County, Maidstone Township, RR at C3, 42°11’ N, 82°46’ W, 10 September 1968, W. Botham 388 TABLE 1—Species new to the Erie Archipelago, Pelee Island, THE CANADIAN FIELD-NATURALIST Vol. 91 and East Sister Island Species Potamogeton crispus Potamogeton foliosus Butomus umbellatus Bromus japonicus Hordeum jubatum Muhlenbergia schreberi Phalaris arundinacea Panicum philadelphicum Panicum dichotomiflorum Cyperus erythrorhizos Carex sartwellii Wolffia punctata Wolffia columbiana Commelina communis Heteranthera dubia Zigadenus glaucus Smilax tamnoides var. hispida Tris pseudacorus Carya ovalis Quercus palustris Quercus velutina Rumex maritimus Cycloloma atriplicifolium Diplotaxis muralis Diplotaxis tenuifolia Lunaria annua Rorippa islandica Sedum sarmentosum Potentilla argentea Rosa palustris Lotus corniculatus Euphorbia dentata Euphorbia cyparissias Euonymus atropurpureus Acer nigrum Lythrum alatum Ludwigia palustris Oenothera parviflora Myriophyllum spicatum Osmorhiza longistylis Ipomea purpurea Hydrophyllum virginianum Myosotis verna Physostegia virginiana Lamium amplexicaule Lamium purpureum Lindernia dubia Linaria dalmatica Viburnum rafinesquianum Lobelia cardinalis Bidens connata Xanthium strumarium Gnaphalium uliginosum Tussilago farfara Tragopogon dubius Erie ~ we Pelee East Sister Archipelago Island am em OK OOK OKO OK OK OOK am KK KK OK ~ KK Pt a ad x KK x A KKK KK KK KK KK OK 821 (CAN); Pelee Island, Stone Road savanna, 15 September 1972, Campbell and Donaldson (WLU); Stone Road, dry open savanna, 7 August 1974, Campbell, Reznicek and Donaldson (CAN, WLU); and E-W road, west end (north side), red cedar savanna glade, 25-26 September 1976, Campbell, Francis and Diebolt (DAO). First collected on the Ontario mainland in 1968, and subsequently on Pelee Island, this prairie species was also collected, as a probable adventive, along a railroad in Dundas (Pringle 1974). Core (1948) recorded it from Kelley’s Island, Ohio, as well. Few plants exist at the Pelee Island sites. Eclipta prostrata (L.) L. (E. alba (L.) Hassk.) Yerba- de-Tago Essex County, Malden Township, east of Amherst- burg, Concession III, 42°16’ N, 83°06’ W, ditch, 12 September 1971, W. Botham 1359 (CAN). Pelee Island, Fish Point, sandy gravelly shore of lagoon, 7 August 1974, Reznicek, Campbell and Donaldson s.n. (CAN, WLU); and Lighthouse Point, new dike, disturbed soil behind J. Harris farm, 4 September 1974, Campbell and Donaldson (CAN). The first Canadian collection of this species was 5 September 1905, on Point Pelee (Klugh 1906), but it was not reported in Canada again until Core (1948) noted it from Pelee Island. Duncan and Stuckey (1970) had recorded it from Lost Ballast Island, Ohio, from where it later disappeared. It is a semi- cosmopolitan, subtropical annual. New Records in the Erie Islands Flora Comparison of our collections with Calvert (1920), Core (1948), Boivin (1953), Stuckey (1968), Duncan and Stuckey (1970), Duncan (1973), and Stuckey (1975) has disclosed that a number of species were new to the Erie Archi- pelago, Pelee Island, or East Sister Island. These are listed in Table 1. The vouchers for all these records are in WLU except for Zigadenus glaucus which is in Herbarium W. Botham and Rosa palustris which is in the Rondeau Provin- cial Park Herbarium. Discussion Although many of the species we have recorded were undoubtedly overlooked by earlier authors, some species have probably spread from the south to Pelee Island recently. Deam (1940) noted at that time that species like Senecio glabellus and Eclipta prostrata were spreading northward. Ammannia coccinea was 1977 reported by Core (1950) and Stuckey (1968) from the north shore of Ohio, and has now reached Pelee Island and Wheatley. Senecio glabellus has spread to some other Erie Islands as well (Stuckey 1975). The recent floods on Pelee Island (1972-73) may well have opened new bare habitat for these species and possibly others like Conobea multifida and Cyperus spp. A number of the rare species reported upon are concentrated in several of the outstanding natural areas left on Pelee Island. Chief among these is the Stone Road savanna (Figure 1), where Chaerophyllum procumbens var. shortii, Eupatorium altissimum, Phacelia purshii, and Spiranthes magnicamporum occur. On the more wooded edges of this savanna occur Triosteum angustifolium, Valerianella intermedia, and V. umbilicata, as well as Ratibida pinnata (Duncan 1973) which occurs elsewhere in southwestern Ontario but rarely. Brown’s Woods or Middle Point (Figure 1) contains a rich assemblage of uncommon species, including Camassia scil- loides, Chaerophyllum procumbens var. pro- cumbens, Corydalis flavula, Euonymus atropur- pureus, and Myosotis macrosperma. Both the Stone Road savanna and Brown’s Woods are threatened with housing develop- ments in the near future. Pelee Island has some of the finest southern plant communities in Canada, and the loss of any one of these would be serious. Fish Point (Figure 1) is one of the most remarkable, and largest, areas of natural wood- land left on Pelee Island. The point has Celtis tenuifolia and Lycopus virginicus as two of its more interesting southern species, together with Camassia_ scilloides, Chenopodium foggii, Hydrophyllum appendiculatum, Morus rubra, and Myosotis macrosperma. The _ so-called “Verbeek savanna’ (Figure 1) on Lighthouse Point contains scattered individuals of Cheno- podium foggii, Corydalis flavula, and Euphor- bia obtusata. There are also uncommon species associated with these, such as Allium cernuum, and a fine stand of Fraxinus quadrangulata, probably the best representation in Canada. The flooding of the north-central portion of Lighthouse Point (Figure 1) has created ‘Lake Henry’ and attendant large areas of drying shore- line. On the drying shores occurs a most interesting early successional flora. Notable CAMPBELL AND REZNICEK: VASCULAR PLANT RECORDS, S.W. ONTARIO 389 species encountered in 1974 and 1975 were Ammannia coccinea, Conobea mutltifida, Eclipta prostrata, Cyperus odoratus var. squar- rosus, and Cyperus erythrorhizos. The last three areas mentioned have been purchased by the Ministry of Natural Resources and the Essex Region Conservation Authority. These are currently designated as Provincial Nature Reserves. We are hopeful that these areas will remain protected from all destructive uses and well buffered from any development. Most of the species mentioned in this article will be placed on the Ontario Endangered Plant lists (G. W. Argus, personal communication 1976). Acknowledgments Assistance in the field was gratefully received from R. Diebolt, G. Donaldson, G. Francis, J. and M. Harris, M. Harris, R. Mitton, P. Pratt, R. Simpson, and D. Perrin, who also prepared the map. Unpublished records were kindly provided by W. Botham (R.R. 1, Cottam) and J. K. Morton (University of Waterloo). Assistance with identification and herbarium records was freely provided by B. Boivin, P. M. Catling, J. M. Laudenbach, J. K. Morton, J. S. Pringle, J. H. Soper, and A. A. Wellwood. T. Duncan (University of Michigan) assisted with some identifications, and made available his unpublished manuscript on flora of the Erie Islands. The hospitality of John and Molly Harris of Scudder, Pelee Island, was most appreciated during our field work on the island; we are greatly indebted. Part of the field work for this paper was carried out under contract with Nature Reserves and Parks Planning of the Ontario Ministry of Natural Resources and Canadian Wildlife Service of Environment Canada, and some of the work was supported by a grant to Campbell from the Canadian National Sportsmen’s Show. Literature Cited Billings, B., Jr. 1862. List of plants observed growing principally within four miles of Prescott, C. W., and for the most part in 1860. Annals of the Botanical Society of Canada 1: 114-140. Boivin, Bernard. 1953. Additions to the flora of the Erie Archipelago (Ontario). Rhodora 55: 224-226. 390 Boivin, Bernard. 1967. Les Celtis du Canada. Naturaliste Canadien 94(5): 621-624. Burgess, T. J. W. 1889. The Lake Erie Shore as a botan- izing ground. Proceedings of the Hamilton Association 1888-9 (Part V): 41-59. Calvert, E. W. 1920. Notes on the fauna and flora of East and Middle Sister and North Harbour Island, Lake Erie. Canadian Field-Naturalist 34: 109-110. Catling, Paul M. 1976. Spiranthes magnicamporum Sheviak, an addition to the orchids of Canada. Canadian Field-Naturalist 90: 467-470. Clausen, Robert T. 1975. Sedum of North America north of the Mexican Plateau. Cornell University Press, Ithaca, New York. Core, Earl L. 1948. The flora of the Erie Islands: An annotated list of vascular plants. Ohio State University, Contribution Number 9. Core, E. L. 1950. The plants of western Lake Erie after fifty years. Center for Lake Erie Area Research and Franz Theodore Stone Laboratory, College of Biological Sciences, Ohio State University. Curtis, John T. 1959. The vegetation of Wisconsin: an ordination of plant communities. University of Wisconsin Press, Madison, Wisconsin. David, R. W. 1976. Nomenclature of the British taxa of the Carex muricata L. aggregate. Watsonia 11: 59-65. David, R. W. and A.O. Chater. 1977. Carex polyphylla Kar. & Kir. and C. lJeersiana Rauschert. Watsonia 11: 253-254. David, R.W. and J.G. Kelcey. 1974. Carex muricata Agg. Botanical Society of the British Isles News 3(1): 15-16. Deam, Charles C. 1940. Flora of Indiana. William B. Burford Printing Company, Indianapolis, Indiana. Dodge, C. K. 1914. Annotated list of flowering plants and ferns of Point Pelee, Ontario and neighbouring districts. Biological Series, Geological Survey, Canada Department of Mines, Memoir 54 (2). Dore, William G. 1961. A centennial floristic census of Prescott, Ontario. Transactions of the Royal Canadian Institute 33: 49-115. Duncan, Thomas. 1973. Three plant species new to Canada on Pelee Island: Triosteum angustifolium L., Valeri- anella umbilicata (Sull.) Wood, and Valerianella inter- media Dyal. Canadian Field-Naturalist 87(3): 261-265. Duncan, Thomas and Ronald L. Stuckey. 1970. Changes in the vascular flora of seven small islands in western Lake Erie. Michigan Botanist 9: 175-200. Fernald, M. L. 1950. Gray’s manual of botany. 8th edition. American Book Company, New York. Gaiser, Lulu O. and Raymond J. Moore. 1966. A survey of the vascular plants of Lambton County, Ontario. Plant Research Institute, Canada Department of Agriculture, Ottawa. Gleason, Henry A. 1952. The new Britton and Brown THE CANADIAN FIELD-NATURALIST Vol. 91 illustrated flora of the northeastern United States and adjacent Canada. New York Botanical Garden, New York. 3 volumes. Gleason, Henry A. and Arthur Cronquist. 1963. Manual of vascular plants of northeastern United States and adjacent Canada. Van Nostrand Company, Princeton, New Jersey. Henderson, Norlan C. 1962. A taxonomic revision of the genus Lycopus (Labiatae). American Midland Naturalist 68(1): 95-138. Jermy, A.C. and 1T.G. Tutin. 1968. British sedges. Botanical Society of the British Isles, London. Klugh, A.B. 1906. Eclipta alba in Canada. Ontario Natural Science Bulletin 2: 46. ; Luer, Carlyle A. 1975. The native orchids of the United States and Canada excluding Florida. New York Botanical Garden, New York. Macoun, John. 1893. Notes on the flora of the Niagara Peninsula and shores of Lake Erie. Journal and Proceed- ings of the Hamilton Association 9: 78-86. Marie-Victorin, Frére. 1935. Flore Laurentienne. Impri- merie de la Salle, Montréal, Québec. Pringle, James S. 1974. Eupatorium altissimum, Tall Thoroughwort, an addition to the flora of Canada. Wood Duck 27(5): 83. Rouleau, Ernest. 1964. Flore Laurentienne. 2nd edition. Les Presses de L’Université de Montréal, Montréal, Québec. Sheviak, C. J. 1973. A new Spiranthes from the grasslands of central North America. Harvard University, Botanical Museum Leaflets 23(7): 285-297. Soper, James H. 1949. The vascular plants of southern Ontario. Department of Botany, University of Toronto and Federation of Ontario Naturalists, Toronto. Soper, J. H. 1962. Some genera of restricted range in the Carolinian flora of Canada. Transactions of the Royal Canadian Institute 34: 1-56. Stuckey, Ronald L. 1968. Aquatic flowering plants new to the Erie Islands. Ohio Journal of Science 68(3): 180-187. Stuckey, Ronald L. 1975. A floristic analysis of the vascular plants of a marshat Perry’s Victory Monument, Lake Erie. Michigan Botanist 14: 144-166. Transeau, E.N. 1935. The prairie peninsula. 16: 423-437. Voss, Edward G. 1972. Michigan flora. Part I: Gym- nosperms and Monocots. Cranbrook Institute of Science and University of Michigan Herbarium, Bloomfield Hills, Michigan, Bulletin 55. Wagner W.H., Jr. 1974. Dwarf Hackberry (Ulmaceae: Celtis tenuifolia) in the Great Lakes region. Michigan Botanist 13: 73-99. Wahl, H.A. 1954. A preliminary study of the genus Chenopodium in North America. Bartonia 27: 1-46. Ecology Received 31 May 1977 Accepted 25 September 1977 Notes Geographic Variation in Dunlins, Calidris alpina, of North America M. RALPH BROWNING National Fish and Wildlife Laboratory, U.S. Fish and Wildlife Service, National Museum of Natural History, Washington, D.C. 20560 Browning, M. Ralph. 1977. Geographic variation in Dunlins, Calidris alpina, of North America. Canadian Field-Naturalist 91(4): 391-393. An evaluation of the geographic variation in Dunlins, Calidris alpina, of North America reveals that the northern Alaska population of the species is distinct from the race of Siberia and that the populations of northeastern Canada deserve nomenclatural recognition. Thus, three races should be recognized as breeding in North America: arcticola Todd, a short- billed race of northern Alaska; pacifica Coues, a longer-billed race of western Alaska; and hudsonia Todd, a race having dark shaft streaks of the undertail coverts, that breeds in northeastern Canada. Key Words: Geographic variation, Calidris alpina, Dunlin, taxonomy, winter plumage. In a recent review of Dunlins, Calidris alpina, MacLean and Holmes (1971) concluded that three races breed in North America. They considered the race breeding in northern Alaska to be sakhalina Vieillot (type locality: Sakhalin Island), a pale-backed and short-billed race that also breeds in northeastern Siberia. The breeding form of western Alaska was considered to be pacifica Coues (type locality: Simiahmoo, Washington), a darker-backed and longer-billed race. MacLean and Holmes also rec- ognized hudsonia Todd (type locality: Southampton Island, Northwest Territories), a race similar to pacifica, as the breeding form of northcentral Canada. The AOU Check-list (1957), however, recognized pacifica as the only race breeding in North America. Separation of the races of C. alpina has been based primarily on the differences in length of culmen and in dorsal coloration. MacLean and Holmes (1971) considered sakhalina to be the breeding race of eastern Siberia and northern Alaska based only on length of culmen since they regarded the differences in dorsal coloration of specimens from the two regions as representing individual variation. MacLean and Holmes (1971, p. 895) stated that their measurements of exposed culmen of specimens from northern Alaska are “indistinguishable” from those of speci- mens from Siberia measured by them and by Kozlova (1962). Measurements made by MacLeanand Holmes revealed that birds from northern Alaska have means in length of culmen that are slightly greater in males, yet slightly smaller in females when compared to their series from Siberia (Table 1). I examined specimens of C. alpina for variation in size and color. There is little variation in wing chord and length of tarsus between North American popula- tions (cf., Todd 1953; MacLean and Holmes 1971); therefore these measurements are not considered further. Measurements of exposed culmen were used to evaluate variation in size. | employed Student’s r- test to demonstrate statistical differences between samples but this was not the only basis for deriving taxonomic conclusions. Although most of my sam- ples are smaller than those of MacLean and Holmes, my samples are sufficiently large for statistical comparisons. I compared measurements of exposed culmen of specimens from Alaska and northeastern Siberia. The means of the exposed culmen of birds from western Alaska are greater than those from northern Alaska (Table 1). Statistical differences between these samples are highly significant for males (¢ = 8.02) and females (t =5.05). Specimens from northern Alaska have means slightly greater than those from northeastern Siberia in males (t= 1.90, P< 0.05) and females (t = 1.37, P<.0.10). These statistical differences be- tween the samples suggest that the close relationship of the northern Alaskan sample with birds from Siberia was oversimplified by MacLean and Holmes, and that especially on the basis of males, the two populations are not indistinguishable as they claimed. I found that the dorsal coloration of specimens from northern Alaska is similar to that of birds from western Alaska and migrants in comparable plumage collected from the west coast of North America in May and early June. Compared with breeding adults collected at Gizhiga in Siberia, the northern Alaska specimens have darker but brighter dorsal coloration. This difference in dorsal coloration between speci- mens from northern Alaska and northeastern Asia has previously been reported by Conover (1945), Todd (1953), and Vaurie (1965). On the basis of dorsal coloration I found the northern Alaskan birds were similar to pacifica, but separable from sakhalina of northeastern Asia. Although I found some overlap in dorsal coloration the population of C. alpina of northern Alaska are characterized as darker-backed 39] 392 TABLE 1—Comparison of the length of exposed culmen (in millimetres) of adult THE CANADIAN FIELD-NATURALIST Vol. 91 Calidris alpina from North America and northeastern Asia Males Females Locality Range n Mean + SD Range n Mean + SD Pt. Barrow, Alaska, breeding: This study 29.8-38.7 56 33.6 + 1.7 31.3-39.8 47 36.4 + 1.8 MacLean & Holmes (1971) 30.5-36.5 25 33.7 34.5-38.5 25 37.0 arcticola (Todd 1953) 33.0-35.0 10 $33),7/ 34.0-39.5 8 37.0 NE Asia, breeding: This study 29.7-35.7 91 32.8 + 1.5 33.8-40.1 12 37.2+ 1.8 MacLean & Holmes (1971) 30.0-37.0 41 32.9 33.0-41.0 20 37.3 sakhalina (Kozlova 1962) 29.0-38.8 16 34.1 33.0-39.8 18 37.1 Western Alaska, breeding: This study 32.5-40.5 Dil 36.8-1.7 35.8-41.4 15 39.1 + 1.8 MacLean & Holmes (1971) 32.0-40.0 55 36.4 37.0-42.0 29 40.1 pacifica (Todd 1953) 35.0-43.0 10 37.4 38.0-43.5 10 41.4 Northern Canada, breeding: This study 34.5-39.3 19 36.1 + 1.2 37.5-40.9 8 39.0-1.2 MacLean & Holmes (1971) 33.0-39.0 23 36.2 37.0-43.0 17 39.4 hudsonia (Todd 1953) 35.0-38.5 10 36.0 37.0-41.0 10 39.0 Western North America, migrants: This study 34.3-41.5 45 Sail se WS) 34.8-44.3 46 40.0 + 2.2 MacLean & Holmes (1971) 34.0-41.0 112 37.2 34.5-45.0 I111 40.5 Northeastern Asia, migrants: This study 32.2-38.8 13 34.8 + 2.1 31.3-40.4 19 BD see a7) MacLean & Holmes (1971) 30.0-36.5 46 3337) 32.5-40.5 33 36.8 than birds from northeastern Asia (sakhalina) and, in exposed culmen, somewhat intermediate between populations from western Alaska and those from northeastern Asia but closer in culmen length to the latter populations. Because of differences in exposed culmen between birds from northern Alaska and western Alaska, Todd (1953) named the northern Alaskan form arcticola, which he characterized as having a shorter exposed culmen than pacifica and darker above than sakhalina from Siberia. I agree with Todd that the birds from northern Alaska that he named arcticola are sufficiently distinct to warrant recognition. Fur- thermore, Holmes (1966a, b, 1970, 1971) has shown that there are differences in breeding density and temporality of breeding behavior between birds from northern and western Alaska. Specimens from northeastern Canada were as- signed to the race hudsonia by Todd (1953) and by MacLean and Holmes (1971). I found the plumage characters ascribed to hudsonia by Todd are not distinct from those of other populations of C. alpina in North America. Moreover, the length of exposed culmen of my sample of Audsonia is similar to that of pacifica (Table 1). As pointed out by Kenneth C. Parkes (personal communication, 1976), however, there are plumage characters that allow separation of hudsonia from pacifica. Parkes described the alter- nate plumage of pacifica as having immaculate white shafts of the undertail coverts. Only very rare individuals of pacifica were said to have one or two feathers with dark hairline shaft streaks. Parkes also found that in hudsonia shaft streaks were present that extended from the flank feathers posteriorly to the black abdominal patch, and in some individuals the streaks include the white lateral upper tail coverts. In contrast, the streaks or spots on the flank feathers of pacifica almost always terminate anterior to the end of the black abdominal patch. According to Parkes, in basic plumage, specimens of hudsonia have gray streaks on the flank feathers that usually extend to the most posterior feather, even when faint, whereas in pacifica, these markings were described as usually confined to the anterior half of the underparts. Thus, as stated by Parkes, specimens of hudsonia and Pacifica may be distinguished even in non-breeding plumages. Specimens at the U.S. National Museum of Natural History completely support Parkes’ observa- tions. I also examined arcticola and sakhalina for streaking and found specimens representing both races to be identical to pacifica. On the basis of these characters I also consider hudsonia a recognizable race. Three races of C. alpina should be recognized as breeding in North America. The race breeding in northern Alaska, arcticola, characterized by its dark TG dorsal color and short culmen, probably migrates to Asia, as suggested by MacLean and Holmes (1971; see also Norton 1971). Birds breeding in western Alaska, pacifica, are longer-billed and dark above, and winter in western North America. Another long-billed race, hudsonia, further characterized by its dark flank streaks and dark shaft streaks of the undertail coverts, breeds in northeastern Canada and migrates south- ward to eastern North America. Acknowledgments I am indebted to the curators of the following museums for the opportunity to examine specimens: American Museum of Natural History, British Col- umbia Provincial Museum, California Academy of Sciences, Carnegie Museum of Natural History, Cleveland Museum of Natural History, Delaware Museum of Natural History, Field Museum of Natural History, Museum of Comparative Zoology, Museum of Vertebrate Zoology, Oregon State Uni- versity Museum of Natural History, University of Alaska, and University of Utah. I am especially grateful to Kenneth C. Parkes who read the manu- script and offered many helpful suggestions. I thank John Farrand, Jr., and Marshall Howe for their encouragement and critical reading of an earlier draft of this paper. Literature Cited American Ornithologists’ Union. 1957. Check-list of North American birds. 5th Edition. Baltimore, Maryland, NOTES 393) 691 pp. Conover, B. 1945. Notes on some North American shore- birds. Condor 47: 211-214. Holmes, R. T. 1966a. Breeding ecology and annual cycle adaptations of the Red-backed Sandpiper (Calidris alpina) in northern Alaska. Condor 68: 3-46. Holmes, R. T. 1966b. Molt cycle of the Red-backed Sand- piper (Calidris alpina) in western North America. Auk 83: 517-533. Holmes, R. T. 1970. Differences in population density, ter- ritoriality, and food supply of Dunlin on arctic and subarctic tundra. Jn Animal populations in relation to their food resources. Edited by A. Watson. Symposium of the British Ecological Society, Aberdeen, Scotland. pp. 303-319. Holmes, R. T. 1971. Latitudinal differences in the breeding and molt schedules of Alaska Red-backed Sandpipers (Calidris alpina). Condor 73: 93-99. Kozlova, E. B. 1962. Fauna of the U.S.S.R.: Birds: Chara- driformes: Limicolae. Volume 2, Section 1, Part 3. (Translation.) MacLean, S.F. and R.T. Holmes. 1971. Bill length, wintering areas, and taxonomy of North American Dunlins, Calidris alpina. Auk 88: 893-901. Norton, D. W. 1971. Two Soviet recoveries of Dunlins banded at Point Barrow, Alaska. Auk 88: 927. Todd, W. E. C. 1953. A taxonomic study of the American Dunlin (Frolia alpina subsp.). Journal of the Washington Academy of Science 43: 85-88. Vaurie, C. 1965. The birds of the Palearctic fauna. Non- Passeriformes. H. F. and G. Witherby, London. 763 pp. Received 18 June 1976 Accepted 31 July 1977 Record of an American Robin Killing a Shrew CHERYL PENNY and RICHARD W. KNAPTON Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Penny, Cheryl and Richard W. Knapton. 1977. Record of an American Robin killing a shrew. Canadian Field-Naturalist 91(4): 393. On 24 October 1976, Penny observed an American Robin (Turdus migratorius) taking a shrew, probably either Sorex cinereus or Microsorex hovyi, as deter- mined from photographs, at the Taiga Biological Station, Wallace Lake, Manitoba. The robin was first noticed chasing the shrew ona grassy area in front ofa cabin. It ran after and pecked the shrew 5 times, lifted it off the ground, and threw it into the air. Eventually the robin picked up the now apparently dead animal, flew to a small knoll, and there proceeded to peck and shake the shrew for a few minutes. Then the bird flew out of sight: it is not known whether the robin flew off with the shrew or whether the shrew was eaten. There are apparently no records of an American Robin actively chasing and killing mammals. L. R. Powers (1973, Condor 75: 248) reports a pair of robins feeding two shrews to their nestlings, and A. C. Bent (1964, United States National Museum Bulletin 196: 50-51) relates an instance of a robin eating a field mouse, but in neither case is it known whether the animals were taken alive or found dead. We thank Spencer G. Sealy for his helpful comments on the manuscript. Received 17 June 1977 Accepted 25 July 1977 394 THE CANADIAN FIELD-NATURALIST Vol. 91 First Record of Anna’s Hummingbird for Alberta HAROLD W. PINEL! and JOHN R. RIDDELL? 11017-19th Avenue N.W., Calgary, Alberta T2M 0Z8 21212 Belavista Crescent S.W., Calgary, Alberta T2V 2Bl Pinel, Harold W. and John R. Riddell. 1977. First record of Anna’s Hummingbird for Alberta. Canadian Field-Naturalist 91(4): 394. On 6 October 1976 an Anna’s Hummingbird (Calypte anna) was observed by the authors in Calgary, Alberta at the home of Kay and M. Mykulak. On 7 October the hummingbird’s song was recorded by the authors, and the bird was sub- sequently photographed by R. Butot and A. Hanners. The tape-recording and photographs are on file at the Provincial Museum of Alberta. The hummingbird remained in the vicinity until 31 October (Kay Mykulak, personal communication). The identifica- tion of the bird as an immature male Anna’s Hummingbird was accepted by the Alberta Orni- thological Records Committee. Initial observations indicated this bird was not one of Alberta’s known hummingbird species because of its large size, habit of singing while perched, and overall color pattern. Closer examination revealed the crimson-violet cap when the bird was in direct sunlight. Further details observed, based on our field notes, include a green nape, back, shoulder patch, and rump with blackish-gray outer tail feathers and primaries. The tail was slightly forked. Under tail coverts were white with five dark gray spots, two pairs ‘immediately posterior to the vent and a fifth most distal spot. The spots became progressively larger posteriorly. Chest and abdomen were white with hori- zontal dark grayish-green spots and streaks concen- trating in the center of the chest to form an apparent breast spot. A white necklace tapering to the nape separated the chest from the throat area. The throat area was white with black streaks and two triangular areas, one under each eye. These black streaks and triangles appeared crimson-violet in direct sunlight. A teardrop-shaped white area was evident behind the eye and a black streak was noted tapering to the bill from the front of the eye. The bill appeared large and slightly decurved. The above observations were made with the use of binoculars and a 20X wide-angle spotting scope. The bird was always very vocal while perched. The bill was held above the horizontal when the bird sang. Its favorite perch was in a lilac shrub (Syringa vulgaris) 1 m off the ground. The bird made frequent visits to artificial bird feeders in the back yard. The first reported sighting of this species in Canada was at Metchosin, British Columbia, in August 1958, and to date there are 131 observations from British Columbia (C. J. Guiguet, personal communication); the most easterly is at Shuswap Lake. Godfrey (1966) acknowledged the sight records from British Col- umbia but listed the species as hypothetical. On 2 January 1968 the first specimen record in Canada was obtained (Guiguet 1968). Since 1971 this species has become recognized as a fairly regular visitor in Alaska (Gibson 1972; Gibson and Byrd 1974, 1975). Zimmerman (1973) considers a 1969 record from Missoula, Montana, as the most surprising in North America, the implication being that its easterly location, almost due south of Calgary, is significant. The Alberta record is the only sighting east of the continental divide north of Texas, and is thus noteworthy in a largely non-migratory species. Zimmerman (1973) notes that most reports of this species outside California are from suburban areas. He postulates that this is a result of a great increase in artificial feeders and an abundance of exotic flowering plants which provide an invaluable source of food during the period when the native flora is dormant. The Alberta record was in a suburban setting richly supplied with exotic plants and artificial feeders. We thank C. J. Guiguet of the British Columbia Provincial Museum and M. T. Myres of the Biology Department, University of Calgary, for assistance in obtaining reference material. Literature Cited Gibson, D. D. 1972. Alaska region. American Birds 26: 105. Gibson, D. D. and G. V. Byrd. 1974. Alaska region. Am- erican Birds 28: 678. Gibson, D. D. and G. V. Byrd. 1975. Alaska region. Am- erican Birds 29: 104. Godfrey, W.E. 1966. The birds of Canada. National Museum of Canada Bulletin 203. 428 pp. Guiguet, C.J. 1968. First specimen record of Anna’s Hummingbird for Canada. Murrelet 49: 9-10. Zimmerman, D. A. 1973. Range expansion of Anna’s Hummingbird. American Birds 27: 827-835. Received 14 January 1977 Accepted 7 August 1977 QT NOTES 395 Le Ver Québécois (Lumbricus festivus) Envahit la Colombie-Britannique JOHN WARREN REYNOLDS Département des ressources foresti¢res, Université du Nouveau-Brunswick, Case postale 4400, Frédéricton, Nouveau- Brunswick E3B 5A3 Reynolds, John W. 1977. Le ver québécois (Lumbricus festivus) enhavit la Colombie-Britannique. Canadian Field- Naturalist 91(4): 395-396. Until now the known North American distribution of Lumbricus festivus, or the Quebec worm, has been restricted to southern Quebec. The species is also present in the provinces of British Columbia, Ontario, New Brunswick, and the state of Vermont. Lumbricus festivus has been found in species associations with most of the lumbricids recorded for Canada. Species associations for L. festivus in North America range from one to six; the most frequent association is with two other species. The genital tumescenes on setae a and b of segments xxix and Xxx are constant for specimens collected in eastern North America but variable for those specimens received from British Columbia. Lumbricus festivus (Savigny, 1826), ou le ver québécois, est un exotique lombric européen signalé pour la premiere fois en Amérique du Nord par Stafford (1902). Ila simplement mentionné, “from Ontario, Quebec, New Brunswick and Nova Scotia” sans renseignement au sujet des habitats ou des endroits. Le deuxi¢me rapport, fondé sur une anté- clitellienne, fut fait par Langmaid (1964) du Comté de Victoria, Nouveau-Brunswick. Dans mes _ récents échantillonnages de lombrics effectués au Nouveau- Brunswick, en Nouvelle-Ecosse, aux Iles-de-la-Made- leine et a l’ We-du-Prince-Edouard (Reynolds 1975a, b, c, 1976a, b), je n’ai pas trouvé de L. festivus. Bien que ne publié que récemment (Reynolds 1977), un échantillonnage en 1972-73 de l’Ontario révéle cette espéce a 10 localités dans sept des comtés de l’est. Des échantillonnages subséquents au Québec (Reynolds 1975d, e, 1976c, données inédites) ont révélé cette espéce a 52 localités dans 35 comtés sur les deux rives du fleuve Saint-Laurent. Pendant cette méme période, une clitellienne de L. festivus fut trouvée a un endroit dans un comté a la frontiére de l’état du Vermont (Reynolds 1976c). Au début de 1977 la répartition connue de L. festivus en Amérique du Nord était centrée dans la vallée du Saint-Laurent et presque exclusivement au sud du Québec. Reynolds (1977, pp. 92-94) a présenté la synonymie détaillée, la diagnose, les tirages longitudinaux ex- ternes, la biologie et la répartition en Amérique du Nord et en Ontario avec une carte de distribution de L. festivus. Quand les protocoles d’échantillonnage sont suivis soigneusement, les espéces de vers de terre se trouvent en association. Le tableau | indique la fréquence du nombre d’espéces associées a L. festivus en Amérique du Nord. Les associations varient considérablement en importance selon lhabitat et la géographie. TABLEAU 1—Fréquence du nombre d’espéces de vers de terre associé a Lumbricus festivus en Amérique du Nord Nombre d’espéces trouvées en association avec Nombre L. festivus de Région localitts 0 1 2 3 4 5 Colombie-Britannique 6 004 1 0 ~0 Ontario 10 OF 223) 5° nll 2220 Québec 52 PU NG Th Ss 2 Nouveau-Brunswick l 00 1 0 0 0 Vermont (E.U.) ] 0 0 0 0 1 +0 Reynolds a révisé (1973; 1977, p. 121) les associations déja établies pour différents habitats. Récemment nous avons recu 120 échantillons de vers de terre du sud de la Colombie-Britannique. Six endroits dans un champ contenaient le L. festivus. C’est la premiere fois que l’on rapporte cette espéce en Amérique du Nord ailleurs que dans la vallée du Saint-Laurent. Le L. rubellus fut également trouve dans tous les six prélévements, Octolasion tyrtaeum dans cing prélévements et Aporrectodea tuberculata dans deux prélévements. Dans les échantillons du Québec, le L. festivus fut trouvé quelques fois en association avec toutes les espéces de vers de terre de la province, sauf Octolasion cyaneum. Le L. festivus fut trouvé en association surtout avec Aporrectodea turgida (45% des 52 prélevements), A//lolobophora chlorotica et Aporrectodea tuberculata (26%), Den- drodrilus rubidus (19%), Dendrobaena_ octaedra (16%) et les huit espéces qui restent dans < 10% des prélevements. Aux dix localités de ’Ontario, le ver québécois fut trouvé sept fois en association avec 396 Echelle | Ss (1H) Kilometres Ficure 1. La répartition connue de Lumbricus festivus en Amérique du Nord. Eisenia rosea, s\x fois avec Aporrectodea tuberculata, trois fois chacun avec Allolobophora chlorotica et Aporrectodea turgida et une fois chacun avec Apor- rectodea trapezoides, Dendrobaena octaedra, Eisen- iella tetraedra et O. tyrtaeum. Un prélévement au Nouveau-Brunswick contenait Aporrectodea tuber- culata et L. terrestris, tandis qwun seul prélévement au Vermont contenait ces deux espéces de méme que Dendrobaena octaedra et L. rubellus. Les soies ventrales sur certains segments dans trois des quatre espéces de Lumbricus rapportées en Amérique du Nord sont sur les papilles puberculien- nes. A lest de Amérique du Nord, L. festivus porte sur les papilles puberculiennes les soies a et b des deux cétés des segments xxix et xxx. Les specimens de L. festivus de Colombie-Britannique portent sur les papilles puberculiennes les soies a et b des deux cétés des segments xxix (un spécimen), Xxix et Xxx (SIX spécimens) et xxx (un spécimen). Un spécimen n’a pas des soies sur les papilles puberculiennes des segments XX1X OU XXX et un autre spécimen porte les soies a et b sur les papilles puberculiennes d’un cété seulement du segment xxix. Les spécimens de L. festivus avec des spermatophores n’ont été trouvés qu’au Québec (Reynolds 1977, p. 92). Ces récents échantillons de la Colombie-Britan- nique créent maintenant un éparpillement plus grand dans la répartition de L. festivus en Amérique du Nord (Figure 1). Il y avait 19 espéces de vers de terre signalées en Ontario (et au Canada) par Reynolds (1977). Les autres provinces de l’est ont été prospec- THE CANADIAN FIELD-NATURALIST Vol. 91 tées méthodiquement: 13 a 16 espéces de vers de terre ont été trouvées dans chacune. Cette mention de L. festivus en Colombie-Britannique augmente la liste provinciale des vers de terre a 14 espéces. Les trois provinces des prairies ont moins de vers de terre inventoriés (Alberta 5, Saskatchewan |, Manitoba 7). Cette situation est probablement due a un manque d’échantillonnage plutdt qu’a une absence des vers de terre. Remerciements L’auteur remercie Valentin Schaefer, Département des sciences biologiques, Université Simon Fraser pour avoir fourni les spécimens de la Colombie- Britannique et William L. Staples, Département des ressources forestiéres, Université du Nouveau-Bruns- wick pour l’exécution de la figure du texte. Il remercie aussi Wilma M. Reynolds et Bernard-M. Thériault pour avoir lu le manuscrit, pour leurs commentaires et leurs suggestions. Litérature Citée Langmaid, K. K. 1964. Some effects of earthworm in- vasion in virgin podzols. Canadian Journal of Soil Science 44: 34-37. Reynolds, J. W. 1973. Earthworm (Annelida, Oligochaeta) ecology and systematics. /n Proceedings of the First Soil Microcommunities Conference. Edited by D. L. Dindal. United States Atomic Energy Commission, National Technical Information Service, United States Department of Commerce, Springfield, Virginia. pp. 95-120. Reynolds, J. W. 1975a. Boiteagan (Oligochaeta: Lumbric- idae) Cheap Breatunn. Megadrilogica 2(6): 1-7. Reynolds, J. W. 1975b. Les lombricidés (Oligochaeta) des Iles-de-la-Madeleine. Megadrilogica 2(3): 1-8. Reynolds, J. W. 1975c. The earthworms of Prince Edward Island (Oligochaeta: Lumbricidae). Megadrilogica 2(7): 4-10. Reynolds, J. W. 1975d. Les lombricidés (Oligochaeta) de la Gaspésie, Québec. Megadrilogica 2(4): 4-9. Reynolds, J. W. 1975e. Les lombricidés (Oligochaeta) de Ve d’Orléans, Québec. Megadrilogica 2(5): 8-11. Reynolds, J. W. 1976a. A preliminary checklist and dis- tribution of the earthworms of New Brunswick. New Brunswick Naturalist 7(2): 16-17. Reynolds, J. W. 1976b. The distribution and ecology of the earthworms of Nova Scotia. Megadrilogica 2(8): 1-7. Reynolds, J. W. 1976c. Catalogue et clé d’identification des lombricidés du Québec. Naturaliste Canadien 103(1): 21-27. Reynolds, J. W. 1977. The earthworms (Lumbricidae and Sparganophilidae) of Ontario. Life Sciences Miscel- laneous Publications, Royal Ontario Museum. x + 141 pp. Stafford, J. 1902. Notes on worms. Zoologischer Anzeiger 25: 481483. Received 15 April 1977 Accepted 17 August 1977 1977 NOTES 397 Melanistic Butler’s Garter Snakes (Thamnophis butleri) at Amherstburg, Ontario P. M. CATLING and W. FREEDMAN Department of Botany, University of Toronto, Toronto, Ontario MSS IAI Catling, P. M. and W. Freedman. 1977. Melanistic Butler’s Garter Snakes (Thamnophis butleri) at Amherstburg, Ontario. Canadian Field-Naturalist 91(4): 397-399 Three melanistic adults and some melanistic young of Thamnophis butleri. are described from an abandoned quarry near Amherstburg, Ontario. This represents the first record of melanism in 7. butleri. Comparison is made with the melanistic phase of Thamnophis sirtalis sirtalis. The ratios of melano to normal in broods from both normal and melanistic females of T. butleri are discussed with respect to the predictions of simple Mendelian genetics, with melanism assumed to be a recessive trait. Key Words: melanism, Thamnophis butleri, Garter Snake, Ontario, recessive trait, genetics. During a study of snake populations in an old limestone quarry approximately 3km north of Amherstburg (42°07’ N, 83°05’ W), Essex County, Ontario, we found three melanistic adult Butler’s Garter Snakes (Thamnophis butleri), which apparent- ly represent the first records of melanism in this species. The first of these melanos, captured on 14 May 1976, is a female with total body length of 54.2 cm; the second, captured on 13 June 1976, is a male 42.2 cm long; the third, captured on 7 May 1977, is a female 34.5 cm long. Of 300 adult Butler Garter Snakes captured, marked, and released in the quarry area during the late spring and summer of 1976 and the early spring of 1977, only these three adults were melanistic. These three snakes are very similar with an overall velvety black appearance above and differ markedly from normal Butler’s Garter Snakes (Figures | and 2). The dorsal and lateral stripes are very dark grayish- olive, becoming lighter prior to shedding. The lower lateral parts and venter are more lustrous and blackish-brown, the brown becoming more apparent toward the lateral stripes. The venter becomes grayish-blue during the pre-shedding dull stage, especially anteriorly. An exception to this overall blackish coloration is the whitish edge of the scales on the chin, beginning with the chin shields and including the first few ventral plates. The only other exceptions include the bright yellow upper half of scale 40 in the third row on the right side of the larger female, and a bright yellow lower half of scale 33 in the fourth row on the left side of the male. Scale counts for two of the three melanistic snakes are summarized in Table |, and fall well within the normal range of 7. butleri. Various characteristic features of 7. butleri are readily apparent in our melanistic specimens, such as the relatively small head, relatively small eyes, prominent brownish FiGurRE |. Normal striped and melanistic 7. butleri from Amherstburg, Ontario. Both snakes are females, and the melanistic individual is 54.2 cm long and |.1 cm across the head. marking on the sides below the lateral stripe, position of the lateral stripe centered on the third scale row, and prehensile behavior when held. The three melanistic adults, as well as some melanistic young resulting from pre-captivity mating of the large female, were kept in captivity for several sheddings. There was absolutely no change in colora- tion. It appears that the melanistic snakes are black from birth and never change their appearance. 398 d FIGURE 2. Detail of snakes shown in Figure |. (a) Anterior lateral view. (b) Ventral aspect at middle of body, melanistic female to the left. (c) Lateral aspect of head, melanistic female above with white edging on chin shields (arrow) distinguishable from glare on upper labials. (d) Anterior dorsal aspect. The melanistic 7. butleri described above differ somewhat from the melanistic phase of the Common Garter Snake (Thamnophis sirtalis sirtalis) as de- scribed from the Lake Erie region by various authors (e.g., Blanchard and Blanchard 1940; Logier 1929, 1958; Froom 1972). Specifically the melanistic T. butleri differ from melanistic 7. sirtalis in having the white on the throat much less well developed, the lateral and dorsal stripes somewhat more conspic- uous, and in having dark brown coloration below the lateral stripes and extending onto the venter. In melanistic 7. sirtalis, the dorsal and lateral stripes are never continuous and never show the normal colora- tion. When present these stripes are limited to a few white or cream-colored scales within 2-5 cm posterior to the head. Also, melanistic 7. sirtalis never show THE CANADIAN FIELD-NATURALIST Vol. 91 normal or even near-normal! coloration below the lateral stripes or on the ventral surface (A. R. Gibson, personal communication). The mating experiments of Blanchard and Blan- chard (1940) suggested that melanism in T. sirtalis from Lake Erie is dependent upon a single Mendelian recessive factor. The large melanistic female gave birth on 14 July 1976 to three normal and six melanistic young. If melanism in 7. butleri is governed by a simple Mendelian recessive factor then we would expecta 1:1 ratio of normal to melanistic young if she had mated with a normally colored but heterozygous male. Mating with homozygous males would result in either all normal or all melanistic young. Our ratio of | normal :2 melanistic offspring may be explained in a number of ways. Blanchard and Blanchard (1940), and more recently Gibson and Falls (1975), were of the opinion that more than one male could be represented in broods where Mendelian ratios are not exact, either through double copula- tions or second matings. It is important to bear in mind, however, that with small samples of offspring, deviations of one or two from Mendelian prediction may not be significant. Five normally colored pregnant females from the Amherstburg quarry were brought into captivity so as to record both the number of broods with melanistic young, and the color phase ratio of the young. All of these females gave birth resulting in broods of 7-11 young, and one brood of 10 contained a melanistic TABLE |. Scutellation of female melanistic Thamnophis butleri from Amherstburg, Ontario Body length (cm) 54.2 34.5 Scale rows 5 rows post. to neck 19 19 Mid-body 19 19 5 rows ant. to cloaca 17 17 Ventral plates 138 137 Caudal scales 59 57 Labials Upper right 6 6 Upper left 6 6 Lower right 7 7 Lower left 7 1 Preoculars Right Left Postoculars Right 2 3 Left 3 3 OMe snake. From the crossing of two heterozygotes, Mendelian genetics would have predicted 2.5 mela- nos. The newborn young in the quarry were surveyed on 10 and 11 July 1976. Although a total of 27 was captured, none of these were melanistic. - The above data may fit the assumption that melanism in the Amherstburg population of 7. butleri is controlled by a simple recessive factor inherited in simple Mendelian fashion, as demonstrated for Lake Erie 7. sirtalis (Blanchard and Blanchard 1940). The possibility of multiple inseminations (Gibson and Falls 1975), however, makes it impossible to base definite conclusions on morph frequencies in off- spring of wild snakes. Rather, crossings will have to be made with captives of known genotype, kept in isolation. All of the melanistic 7. butleri from Amherstburg were released there later, excepting for the large adult female which died in captivity and was placed in the collection of the National Museum of National Sciences, Ottawa (No. 17686). In addition to this specimen, there are photographs on file at the National Museum and the Royal Ontario Museum, Toronto. NOTES 399 Acknowledgments We gratefully acknowledge Francis Cook of the National Museum of Natural Sciences, Ottawa: Craig Campbell of Waterloo, Ontario; Janet Planck of Kitchener, Ontario; and Roger Conant of the Univer- sity of New Mexico for their critical reading and helpful suggestions. Literature Cited Blanchard, F.N. and F. C. Blanchard. 1940. The inheri- tance of melanism in the Garter Snake (Thamnophis sirtalis sirtalis (Linnaeus)) and some evidence of effective autumn mating. Michigan Academy of Science, Arts and Letters, Papers 26: 177-192. Froom, B. 1972. The snakes of Canada. McLelland and Stewart Ltd., Toronto. 128 pp. Gibson, A. R. and J. B. Falls. 1975. Evidence for multiple insemination in the Common Garter Snake, Thamnophis sirtalis. Canadian Journal of Zoology 53(9): 1362-1368. Logier, E. B.S. 1929. Melanism in the Garter Snake, Thamnophis s. sirtalis, in Ontario. Copeia 172: 83-84. Logier, E. B.S. 1958. The snakes of Ontario. University of Toronto Press, Toronto. 94 pp. Received 19 May 1977 Accepted 18 August 1977 An Improved Design for a Small Mammal Live Trap ANDREW RADVANYI Canadian Wildlife Service, Fisheries and Environment Canada, 10025 Jasper Avenue, Edmonton, Alberta TSJ 1S6 Radvanyi, Andrew. 1977. An improved design for a small mammal live trap. Canadian Field-Naturalist 91(4): 399-401. Inclusion of variable quantities of bait and nesting materials have frequently altered the sensitivity and effectiveness of small mammal live traps. An improved design of a light weight small mammal live trap is described in which the extreme sensitivity is unaffected by variations in quantity of either bait or nesting material placed inside. Key Words: small mammal live trap, improved design, trap sensitivity, effectiveness. Numerous articles published in past years have described small mammal populations in a diversity of habitats. A wide spectrum of techniques and field equipment used in obtaining the information render as questionable comparisons of trapping ‘results. In describing small mammal populations, major em- phasis is usually placed upon species present, numbers of animals handled, and floristic habitat. Only minor reference is usually made to field equipment used to obtain the data or the techniques employed. Wiener and Smith (1972) compared the relative) efficiencies of four types of small mammal traps in! common use. Similar comparisons have been noted by Neal and Cock (1969), Edwards (1952), Smith et al. (1971), Duran (1968), and Goodnight and K oestner (1942). In small mammal studies over the past 17 years I have made extensive use of sheet-metal traps based on the Sherman live trap design. Although particular attention was repeatedly given to adjustment of the traps to ensure sufficient sensitivity to enable capture of even the smallest and most light-weight animals, several disadvantages inherent in the features of the traps became apparent and led to the design of an improved small mammal live trap outlined here. Evans (1975) and Wiener and Smith (1972) em- phasized the importance of adjusting trap sensitivity prior to use. Unfortunately the sensitivity can be 400 THE CANADIAN FIELD-NATURALIST Vol. 91 Xx Treadle IS cm long Bait holder Wire holder for nest material 9 cm Spring Front door in set position FiGureE |. Cutaway schematic drawing of the improved small mammal live trap, shown in the set position. altered by the amount of bait and nest material placed inside the trap, Application of peanut butter and oatmeal bait smeared onto the inside of the rear door (Wiener and Smith 1972) provides a partial solution to the trap sensitivity problem but limits the quantity of bait used in the trap. Sufficient food and nesting material must be included to sustain the captive animal until the traps are checked. The problem of trap sensitivity being affected by varying weight of bait and nesting material appeared to have been solved in the live trap designed by Buech (1974). The Buech trap, although extremely sensitive to animal weight, cannot be set without first removing the lid. To set the trip mechanism, replace the lid, fasten the latter to the trap by means of two elastic bands, and then put the trap into place without springing it, requires extreme care. The modifications incorporated in the new trap design are aimed at improving the shortcomings mentioned. The trap dimensions (Figure 1) can be altered to accommodate prey species. Although non- collapsible, the traps are of light-weight 0.030-guage aluminum and weigh only approximately 206 g — an important consideration when large numbers of traps must be packed into a remote study location. Theoret- ically the traps could be made collapsible by inclusion of appropriate interlocking piano hinge mechanisms along the long edges of the trap body, but to do so would increase appreciably the construction cost per trap. The overlapping front door-—treadle release mechanism is extremely sensitive. A penny weight (3.17 g) placed gently upon the distal end of the treadle is amply sufficient to spring the trap. The smallest of mammals we encountered in our field studies (i.e., Sorex cinereus) weigh 2.5 to 4.0 gand are readily captured in the traps. The new design lacks the brace plate extending across the proximal end of the treadle as found in the Sherman trap. Mice could conceivably perch on this plate and remove the bait beyond without activating the treadle release mech- anism. In our trapping procedure a large table- spoonful of a paste bait and a small slice of apple are placed into the bait holder inside the back door of the trap. As with the bait, the nesting material provided in 1977 the trap does not contribute any additional weight onto the treadle release mechanism. The nesting material is inserted behind a short hooked wire fastened to a rivet in the trap ceiling. The sensitivity of the trap is adjusted by the curvature put on the spring wires under the front door and treadle. The greater the bend in the ends of the wires, the greater the pressure exerted on the door and treadle while the trap is in the set position. Unlike the Buech live trap, our trap can easily and quickly be reset from the closed position by inserting one hand inside the trap, pressing down on the front door and treadle until the former catches under the latter, and then withdrawing the hand — a 3- to 4-s Operation. Of 550 small mammals handled in 1691 captures and recaptures during our 1976 field season, trap casualties among captured mice amounted to only 2.7%, even though climatic conditions ranged from hot days to traps buried under 25.4-30.5 cm snow. A three-sided plywood and lumber sheltering device placed over each trap served to minimize animal losses due to temperature extremes and inadvertent spring- ing of the sensitive traps by wind, rain, or hail. NOTES 401 Literature Cited Buech, R. R. 1974. A new live-trap and techniques for winter trapping small mammals. Canadian Field- Naturalist 88(3): 317-321. Duran, J.C. 1968. Comparison of live traps with snap traps. Journal of the Arizona Academy of Science 5: 18. Edwards, R. Y. 1952. Howefficient are snap traps in taking small mammals? Journal of Mammalogy 33: 497-498. Evans, B. A. 1975. Sensitizing Sherman small mammal live traps to improve their efficiency. Northwest Science 49(3): 160-162. Goodnight, D. J.and E. J. Koestner. 1942. Comparison of trapping methods in an Illinois prairie. Journal of Mammalogy 23: 435-438. Neal, B. R. and A. G. Cock. 1969. An analysis of the selec- tion of small African mammals by two break-back traps. Journal of Zoology 158: 335-340. Smith G. C., D. W. Kaufman, R. M. Jones, J. B. Gentry, and M.H. Smith. 1971. The relative effectiveness of two types of snap traps. Acta Thierologica 16: 240-244. Wiener, J. G. and M. H. Smith. 1972. Relative efficiencies of four small mammal traps. Journal of Mammalogy 53(4): 868-873. Received 17 March 1977 Accepted 10 August 1977 Collections of Spiders beneath Snow J. OLYNYK and R. FREITAG Department of Biology, Lakehead University, Thunder Bay, Ontario P7B 5El Olynyk, J. and R. Freitag. 1977. Collections of spiders beneath snow. Canadian Field-Naturalist 91(4): 401-402. Observations of activities of invertebrates in snow environments have been recorded by relatively few workers. Novikov (1940), Chapman (1954), Wolska (1957), Strubing (1958), Kuhnelt (1961), Kevan (1962), Nasmark (1964), Fjellberg and Greve (1968), Hagvar (1971, 1973), and Aitchison (1974) have found that a variety of invertebrates including insects and spiders are active on the snow surface as well as beneath the snow. Spiders were collected throughout the winter of 1976-77 as part of a year-long ecological study on carabid beetles and their associated mite populations near Thunder Bay, Ontario. During the winter no beetles, but 13 spiders, were captured. Two collecting sites were established in mixed spruce forest. One was beside Highway 11, 0.8 km from the junction of Highways 11 and 17, 65 km west of Thunder Bay and the other beside Gilbride Road near One Island Lake 32 km northwest of Thunder Bay. On each site were placed 100 dry pitfall traps 7 cm deep and 11.5 cm in diameter at the top. A piece of 1.5-cm-mesh chicken-wire, 15 cm square, covered each trap preventing the entrance of larger animals. Aluminum pie plates over each trap prevented the entrance of precipitation and debris. The last pre-winter collection was made on 9 November, at which time patches of snow were present. After that date the traps were considered to be snowed-in for the winter. Activity was considered to be unlikely on the soil surface until a subnivean environment was established. Collections were made from 25 traps in each site early in each of the months January through April inclusive. By collecting from different traps each time, disturbance of snow cover was eliminated as a factor. On 6 April, all but 16 of the 50 traps checked were embedded in a layer of solid ice beneath a thin covering of snow. Nine of those 16 traps were also full of ice. The seven traps free of ice contained no specimens. Thirteen spiders representing four species and three 402 THE CANADIAN FIELD-NATURALIST TABLE 1—Spiders collected in winter 1977 beneath snow near Thunder Bay, Ontario Number Snow Date Taxa and sex depth (cm) 6 Jan. Clubionidae: Agroeca ornata ] ie) 25 Linyphiidae: Lepthyphantes zebra panier 45 Amaurobiidae: Callioplus euoplus 3 @ 45 Amaurobius borealis 1 imm. 45 8 Feb. Amaurobius borealis 2 Q 45 7 Mar. Amaurobius borealis 1 Q 30 Amaurobius borealis 1 fe) 50 6 Apr. == Snow patches families were collected over the winter (Table 1). Nine of these spiders were either active when first observed in the traps or became active before they were placed in 70% ethanol in the field. The spiders captured are common leaf-litter inhabitants of boreal forests. We thank R. E. Leech for the spider identifications, B. Barnes for technical assistance, and the National Vol. 91 Research Council of Canada for financial support (Grant No. A4888). Literature Cited Aitchison, C. W. 1974. A technique for sampling active subnivean invertebrates in Southern Manitoba. Manitoba Entomologist 8: 32-36. Chapman, J. 1954. Observations of snow insects in western Montana. Canadian Entomologist 86: 357-363. Fjellberg, A. and L. Greve. 1968. Notes on the genus Boreus in Norway. Norsk Entomologisk Tiddskrift 15(1): 33-34. Hagvar, S. 1971. Field observations on the ecology of a snow insect, Chionea araneoides Dalm. (Diptera: Tipu- lidae). Norsk Entomologisk Tiddskrift 18: 33-37. Hagvar, S. 1973. Ecological studies on a winter-active spider Bolyphantes index Thorell (Araneida: Linyphi- idae). Norsk Entomologisk Tiddskrift 20: 309-314. Kevan, D. K. McE. 1962. Soil animals. H. F. & G. Wither- by, London. 237 pp. Kuhnelt, W. 1961. Soil biology with special reference to the animal kingdom. Faber & Faber, London. 379 pp. Nasmark, O. 1964. Vinteraktivitet under snou hos land- levande evertebrater. Zoologisk Revy 26: 5-15. Novikov, G. 1940. Observations on insects on the snow at the Arctic Circle. Priroda (Moskva) 3: 78. (In Russian.) Strubing, H. 1958. Schneeinsekten. Die neue Brehm- Bucheri. A. Ziemsen Verlag, Wittenberg, Lutherstadt. 47 pp. Wolska, H. 1957. Preliminary investigations on the thermic preferendum of some insects and spiders encountered in snow. Folia Biologica (Krakow) 5: 195-208. Received 6 June 1977 Accepted 25 August 1977 Northern Range Extension for the Brassy Minnow in Northeastern Alberta DAVID K. BERRY Fish and Wildlife Division, Alberta Department of Recreation, Parks and Wildlife, Edmonton, Alberta T6H 4P2 Berry, David K. 1977. Northern range extension for the Brassy Minnow in nertheastern Alberta. Canadian Field- Naturalist 91(4): 402-403. In 1976 intensive sampling of the Athabasca River north of Fort McMurray provided several specimens identified as Brassy Minnow, Hybognathus hank- insoni. Previous records for this species are from the Milk River drainage in southeastern Alberta (Paetz and Nelson 1970), Musreau Lake of the Peace River drainage in northwestern Alberta (Bishop 1975), anda single specimen from the Athabasca system men- tioned in Bishop (1975). This specimen was identified from collections made by W. E. Griffiths and D. B. Ferster (1973. Preliminary fisheries survey of the Winefred Pelican Area. Alberta Department of Lands and Forests, Fish and Wildlife Division, Survey Report Series 19. 449 pp.), while they were surveying the upper House River (55°50’N, 112°20’W) south of Fort McMurray. The Brassy Minnow occurs from the Upper St. Lawrence River and Lake Champlain region of New York, west through southern Ontario and Michigan, west through the Arkansas and Missouri Rivers to Colorado, Wyoming, and Mon- tana, north to Alberta, and in British Columbia in the lower Fraser River and headwaters of the Peace River Ia (Scott and Crossman 1973). Nineteen specimens were collected in beach seines at three locations in the Athabasca River. On 19 July 1976, 14 Brassy Minnows were taken at the confluence of the Horse River (56°44’N, 111°28’W). In Sep- tember four specimens were found in subsamples from island sites near Fort McMurray, while one specimen was collected 28 km further down river. In this region the Athabasca is a wide, slow, and highly turbid river. Most of the tributaries in the area are low- gradient brown-water steams. Measurements made on the preserved sample were as follows: total length 49 to 72 mm; fork length 44 to 66 mm; lateral line scales 37 to 41 (mode 39); scales above lateral line six; anal and dorsal fin rays eight; scale radii 13 to 19. Four specimens have been placed in the Museum of Zoology, University of Alberta, Edmonton (UAMZ #3766). McPhail and Lindsey (1970) hypothesized that a postglacial southeast drainage route existed, allowing the dispersal of fish from the Missouri system into northern waters; and predicted that the Brassy Minnow may be discovered in some additional Alberta or Saskatchewan waters that lie along the supposed dispersal route. Bishop (1975) supported the existence of such a route. The presence of Brassy NOTES 403 Minnow in the Athabasca drainage near Fort McMurray gives further support to the hypothesis and extends the known range northward in Alberta 510 km. My thanks to Wayne Roberts, Museum of Zoology, University of Alberta, for his confirmation of my identification. The collection of Brassy Minnows from the Athabasca River was incidental to a larger study with funding from the Aquatic Fauna Com- mittee of the Alberta Oil Sands Environmental Research Program. Literature Cited Bishop, F. G. 1975. A new distribution record for brassy minnow in northwestern Alberta. Canadian Field- Naturalist 89: 319-320. McPhail, J. D. and C. C. Lindsey. 1970. Freshwater fishes of northwestern Canada and Alaska. Bulletin of the Fisheries Research Board of Canada Number 173. 381 pp. Paetz, M. J. and J. S. Nelson. 1970. The fishes of Alberta. Government of Alberta, Queen’s Printer, Edmonton. 282 pp. Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes of Canada. Bulletin of the Fisheries Research Board of Canada Number 184. 966 pp. Received 11 July 1977 Accepted 21 August 1977 Distribution of the Auricled Twayblade Orchid (Listera auriculata) in Canada and Description of New Stations in Southern Ontario R. EMERSON WHITING! and PAUL M. CATLING2 174 Denison Road West, Weston, Ontario MIN 1C2 2Department of Botany, University of Toronto, Toronto, Ontario M5S IAI Whiting, R. Emerson and Paul M. Catling. 1977. Distribution of the auricled twayblade orchid (Listera auriculata) in Canada and description of new stations in southern Ontario. Canadian Field-Naturalist 91(4): 403-406. On the basis of herbarium specimens overlooked by recent authors, the range of Listera auriculata Wiegand (Orchidaceae) is extended northwestward into Manitoba and northward in Ontario, Quebec, and Newfoundland. The known distribution in Ontario is also extended southward by recent discoveries at five locations in Hastings and Haliburton Counties. The newly discovered stations are described in terms of associated plant species and soil factors. Distribution maps for Canada and for southern Ontario are included. Key Words: orchid, Listera auriculata, distribution, range, southern Ontario. The auricled twayblade, Listera auriculata Wie- gand, was one of the last species of the genus Listerato be described in North America (Wiegand 1899). Its distribution and habitat have not always been adequately reported, for the species is rare over much of its range and has been confused with other Listera species. Several recent botanical works, including those of Correll (1950), Fernald (1950), Gleason and Cronquist (1963), and Luer (1975), give a more restricted distribution in Canada than the species actually has. The following collections indicate the presence of L. auriculata at locations, plotted in Figure |, that are about 780 km northwest and 710 km north of the area of distribution shown by Luer (1975, p. 87). Manitoba: Duck Mountain Provincial Park, Singush Lake, 51°36’ N, 100°48’ W, 21 July 1954, J. S. Rowe 404 THE CANADIAN FIELD-NATURALIST FiGuRE 1. Canadian distribution of Listera auriculata, previous publication. 668 (DAO). This is the northwestern limit of presently known distribution. Ontario: Kenora District, Big Trout Lake near Trout Lake Hudsons Bay Company post, 53°49’N, 89°53’ W, 31 July 1956, 1. Hustich 1630 (CAN). Quebec: New Quebec Territory, Poste-de-la-Baleine (Great Whale River), Baie d’Hudson, ca. 55°17’N, 77°46’'W, 4 August 1969, S. Brisson et P. Forest (LKHD, SFS); New Quebec Territory, Fort Chimo area, Highfall Creek, 58°01’N, 68°29’W, 2 August 1967, Y. Makinen 67-1584 (sub L. convallarioides CAN). This specimen represents the northern limit of known distribution. Newfoundland: Labrador West District, Hamilton River west of Lake Winikapau, 27 June 1967, J. Hustich 127 (CAN); Labrador North District, northeast side of Grand Lake, 9 July 1967, I. Hustich 299 (CAN); Labrador North District, Northwest River area, 9 July 1963, 7. Hustich and P. Kallio 321 (CAN); Labrador North District, Goose Bay, Hamilton River, 53°19’ N, 60°21’ W, 13 August 1949, W. B. Schofield (sub L. cordata, DAO). Vol. 91 Listera quriculata Wieg. dots (@) based on examination of specimens at CAFB, CAN, DAO, LKHD, PMAE, SFS, UAC and WIN. *, distribution limits after * Luer (1975) SCALE ° 100 based on recently examined herbarium specimens and The Canadian distribution map (Figure 1!) suggests that the greatest frequency of occurrence is close to the shores of Lake Superior, but it should be remembered that these shores have been more extensively botanized than have areas to the west, north, and east. In northern Ontario there are only three adequately documented stations that are far from Lake Superior. In southern Ontario the auricled twayblade was first discovered in 1968, and at present we know of seven widely separated locations (Figure 2). Two of these have already been described: Renfrew County near: Barry’s Bay (MacKenzie and Greenwood 1969) and: Nipissing District, Bronson Township (Brunton and Crins 1975). The other five stations are described below. 1. Haliburton County: Snowdon Township, 8 km northeast of Kinmount, 44°50’ N, 78°34’ W. Here, on 12 July 1974, we discovered two colonies of three and eight plants under conifers (Abies balsamea) and alders (A/nus rugosa), on sandy banks of the Irondale River (R. E. Whiting, TRT). This appears to be the southernmost station of L. auriculata in Canada, extending the known range about 75 km south- westward. 1977 KILOMETRE Ontario, based on examination of specimens at CAN, DAO, LKHD, and TRT. 2. Hastings County: Faraday Township, 3 km north of Bancroft, 45°05’ N, 77°52’ W. Here, on 13 July 1974, we discovered about 50 plants rooted in acid sand of a seasonally flooded bank of the York River (P. M. Catling, R. E. Whiting and S. M. McKay, TRT). In addition to the prevalent alders, frequent associates were the moss Atrichum crispum, the liverwort Pellia epiphylla, and a sedge, Carex intumescens. Also present were Picea glauca, Acer rubrum, Ilex verticillata, and Viburnum cassinoides. 3. Hastings County: Dungannon Township, 10 km east of Bancroft, 45°04’ N, 77°44’ W. On 13 July 1974 we saw 15 plants in loose sand of alder thickets along the east bank of the York River(P. M. Catling, R. E. Whiting and S. M. McKay, TRT). 4. Hastings County: Wicklow Township, near Maple Leaf, about 45°17’N, 77°49’ W. Listera auriculata may be seen in at least three places in the Maple Leaf area. (a) We were directed to this vicinity by L. G. Roberts (personal communication) who, on 3 July 1974, had seen 11 plants under alders beside Papineau Creek, 0.8 km north-northeast of Maple Leaf. Here, on 6 July 1976, we found about 60 shaded plants in acid sandy soil (pH 4.7-4.8) that had apparently been NOTES 405 deposited during spring flooding (P. M. Catling and R. E. Whiting, TRT). (b) Farther downstream on Papineau Creek, about 2.5 km east of Maple Leaf and upstream from the site of an old sawmill, we found two colonies of three and nine plants (P. M. Catling and R. E. Whiting, TRT). Here the plants were growing in black muck of alder thickets at the base of a slope. Although the soil was composed of humus instead of sand, it was acid (pH 4.5) and probably subject to flooding during the early spring thaw. Prominent associates (in addition to the prevalent alder) included Onoclea sensibilis, Athyrium filix-femina, Carex intumescens, Maianthemum canadense, Rubus pubescens, and Viola sp. (c) On Little Papineau Creek, about 4 km north-northeast of Maple Leaf, we discovered another colony of about 100 plants in acid sandy soil (pH 4.1-4.2) in periodically flooded alder thickets (P. M. Catling and R. E. Whiting, TRT). In addition to the associates listed above for station 2 on the York River, we saw here Equisetum pratense, Onoclea sensibilis, Athyrium filix-femina, Glyceria striata, Carex crinita, Carex gracillima, Thalictrum poly- gamum, Rubus pubescens, Hypericum ellipticum, Viola sp., Scutellaria lateriflora, and Viburnum trilobum. 5. Hastings County: Mayo Township, near the Little Mississippi River 0.6km_ southeast of McArthurs Mills, 45°07’ N, 77°34’ W. Here, under alders near the foot of a north-facing slope that was covered with hemlock (7suga canadensis) and fir (Abies balsamea), we found a colony of 25 plants (P. M. Catling and R. E. Whiting, TRT). The soil was composed of a black mucky humus (pH 5.3-5.8) and was apparently subject to early spring inundation. The habitat appeared to differ from that of the sawmill stations (4(6) above) chiefly by the presence of a few additional associates: Thalictrum polygamum, Platanthera psycodes, Scutellaria lateriflora, and the moss Climacium dendroides. These new southern Ontario stations are similar to those we have seen in northern Ontario. At Perry Bay, on the Sibley Peninsula in Thunder Bay District, L. auriculata was found in mixed alder and cedar (Thuja occidentalis) swampland and was rooted in acid cushions of sphagnum moss and alder litter. Sandy substrate habitats of L. auriculata are well known along the Lake Superior shores, in both Ontario and Michigan, and along some of the major rivers draining into the lake (Case 1964a, b; Catling 1976). We have found the auricled twayblade in both humic. and sandy substrates, but the soil reaction has always been acid. Alders have always been present, and spring flooding has been evident, sometimes from the 406 presence of debris caught in the lower branches of the alders. It is likely that more stations of L. auriculata will be discovered in Canada. With its preference for moist, acid, sandy or humic substrates and a cool micro- climate, however, it seems unlikely that this species will be found much farther to the south in Ontario. The auricled twayblade blooms during late June and early July in southern Ontario, and until late July farther north. We are grateful to the officials of the various herbaria mentioned in Figures | and 2, to L.G. Roberts for directing us to the first Maple Leaf site, to S. M. McKay for field assistance in the Bancroft area, and to A. A. Reznicek for critical reading of our script. Literature Cited Brunton, D. F. and W. J. Crins. 1975. The auricled tway- blade (Listera auriculata) orchid in Algonquin Park, Ontario. Canadian Field-Naturalist 89(2): 189-190. Case, F.W. 1964a. A hybrid twayblade and its rarer parent, Listera auriculata, in northern Michigan. Michi- THE CANADIAN FIELD-NATURALIST Vol. 91 gan Botanist 3(2): 67-70. Case, F. W. 1964b. Orchids of the western Great Lakes region. Cranbrook Institute of Science Bulletin 48. 147 pp. Catling, P.M. 1976. On the geographical distribution, ecology and distinctive features of Listera Xveltmanii Case. Rhodora 78(814): 261-269. Correll, D.S. 1950. Native orchids of North America. Chronica Botanica Company, Waltham, Massachusetts. 399 pp. Fernald, M. L. 1950. Gray’s manual of botany. 8th edition. American Book Company, New York. 1632 pp. Gleason, H.A. and A. Cronquist. 1963. Manual of vascular plants of northeastern United States and adjacent Canada. Van Nostrand Reinhold Company, New York. 810 pp. Luer, C. A. 1975. The native orchids of the United States and Canada. New York Botanical Garden. 361 pp. MacKenzie, H.N. and E. W. Greenwood. 1969. Range extensions of Listera auriculata Wiegand in Ontario and Quebec. Canadian Field-Naturalist 83(1): 55-56. Wiegand, K.M. 1899. A revision of the genus Listera. Bulletin of the Torrey Botanical Club 26(4): 157-171. Received 17 March 1977 Accepted 25 July 1977 Thermal Selection and Related Behavior in Larval Yellow Perch (Perca flavescens) JAY Ross, P. M. POWLES, and MICHAEL BERRILL Department of Biology, Trent University, Peterborough, Ontario K9J 7B8 Ross, Jay, P. M. Powles, and Michael Berrill. 1977. Thermal selection and related behavior in larval yellow perch (Perca flavescens). Canadian Field-Naturalist 91(4): 406-410. Thermal preferenda and behavior of larval yellow perch (Perca flavescens) were observed at successive developmental stages in a vertical thermal gradient. Newly hatched larvae acclimated to 20, 23, and 25°C selected 24.3+3.9, 242+4.5, and 21.7 + 2.4°C respectively. Preferenda did not differ significantly with development stage to day 24. Prior to swim bladder inflation, equilibrium required a combination of sinking and locomotory activity. After inflation (7-10 days from hatching at 23°C), vertical position was regulated by bladder control and blundering into warm water was reduced. This maladaptive. behavior in response to elevated temperatures (rising to very warm surface layers) is described and discussed. Key Words: Perca, perch, larvae, thermal, temperature, selection, preferenda, shock, behavior. The increasing rate of appearance of once-through cooling systems for electric power generators has emphasized the need for research dealing with the thermal preferenda and behavior of aquatic organ- isms. The early life stages of fish have been largely neglected in this respect, though it has been noted that the young of many species have higher preferenda than older individuals (Fry 1937). Lethal effects (LTs0s) have been studied for only a few species (see Hokanson and Kleiner 1974; Powles 1974). Work on thermoregulatory behavior (thermal selection) in larvae is almost entirely lacking (but see Evero- peyzera 1974; Hoagman 1974). A comparison of the data of F. D. McCracken and S. H. Starckman (1948. Preliminary observations on the preferred temperature of the perch. MS, Ontario Fisheries Research Library, University of Toronto. 4 pp.) for adult yellow perch (Perca flavescens) with that of Ferguson (1958) for yearlings, indicates that adults have cooler preferenda. McCauley and Reid (1973) have verified this observation. In his work on yellow perch, Houde (1969) noted that although larvae under 6.5 mm were able to swim actively, only larger individuals exhibited rheotaxis. In this study we 1977 investigate the possibility that behavior in a vertical thermal gradient shows changes similarly related to development. Our results are the first published observations for larval yellow perch. Our choice of species was further influenced by its value as a food and sport fish and by its almost ubiquitous occurrence in temperate North America. Also, yellow perch spawn littorally and hatch in shallow water close to shore. Exposure to steep thermal gradients similar to those devised in our laboratory are therefore especially likely to be encountered by this species in thermal discharge situations. Methods Thermal selection experiments were conducted ina vertical length of clear plexiglass tubing 70 cm tall and 22cm in diameter. The wall of this tube was horizontally scored into ten 7-cm sections with a mercury thermometer mounted through the wall of each section. Heat was provided by a painted copper coil (5 mm diameter) through which hot tap water could be made to flow from top to bottom. A valve at the bottom permitted introduction of river water from a refrigerated head tank and a similar valve at the top was used to carry away overflow. Two complete perch egg strands were collected on 13 May 1976 from Clear Lake, near Peterborough, Ontario. Eggs were transported to the laboratory, acclimated to an incubation temperature of 25°C; and aquaria at three different temperatures. One strand had reached an advanced developmental stage (embryos moving, eyes pigmented) when collected. This group (A) was divided into two subsamples: A,, acclimated to an incubation temperature of 25°C; and A,, which was allowed to approach room temperature (23°C). These temperatures remained constant (+ 0.5°C) throughout the experimental period. Mass hatch occurred overnight 14-15 May (29-43 h after collection) for Group A; and on 16 May (46-50 h after collection) for group Ao. The second strand (group B), which were inactive and the eyes unpigmented, were placed in a constant- temperature bath adjusted to 20°C. Mass hatching occurred overnight May 16-17 (78-91 h after col- lection). Larvae were fed daily with fresh plankton from the spawning grounds. No attempt was made to estimate or regulate the types and amounts of food provided. Tank hygiene was maintained by siphoning off suspended material daily. A schedule for the thermal selection experiments is given in Table 1. Group A) (23°) was studied throughout its development, until 24 days old, at which time the experiments were terminated because of a shortage of specimens. Experiments with Groups NOTES 407 TABLE 1—Mean and modal temperatures selected by larval yellow perch, Perca flavescens, acclimated to different temperatures at various stages after hatching (May 1976, Clear Lake, near Peterborough, Ontario) Selected Temp. (°C) Acclimation Perch temp., °C N Runs age Mode Mean SD 25 (A,) 31 1] 12h 21.0 PA Vd Pa} 21 5 Dy” 20.5 PBI) Bo 24 6 3h) 2 21.0 DIAS 23 19 8 60” 22.5 WAS Bx? 23 (A,) 37 7 12h 24.0 24.2 4.5 28 8 a3) 7 24.5 24.1 3.8 23 7 3 7 DD.) TMEV Slos) 22 9 47” Des) BND: By) 28 6 3 days DEES 23.5 35 DS) 7 4% DOES Mos) BoP 19 9 Sie 24.0 WB BY 23 8 ne DES) 7S) 3) 18 5 NOs 23.5 Det = ST 19 6 37 Dies Mey Bail 18 9 Ny? Des) 244 44 23 7 Ky) 25.0 248 3.9 21 9 24 ” 24.5 240 2.8 20 (B) 29 6 12h 22.0 DAS 39 24 6 Sil 7 25.0 24.1 3.1 Di 6 43” 21.0 DD = 2D 24 5 60” 21.0 Wp 33,33 25 8 Ta? Piles PSS DT] A, and B were continued only for the first 3 days after hatching. For each “run,” the surface of the gradient was brought to a temperature approximately equal (+ 1°) to acclimation temperature and 20 to 30 larvae were introduced. The gradient was then adjusted to provide an adequate range of temperature and 30 min was allowed for familiarization. The temperature profile was recorded to the nearest 0.5°C and the larvae occurring in each cell were enumerated. Larval counts were performed 3 times in succession (top to bottom) for each temperature profile. A number of runs were performed at 5- to 10-min intervals, the position of preferendum changed, and several more runs were performed. This constituted an “experiment.” Data were then integrated to produce a single temperature distribution by summing the cell counts in each run and, assuming a linear gradient between adjacent cells, assigning these sums to each 0.5°C interval represented. This produced a continuous temperature distribution for each run. Data were then summed for the entire experiment and resultant distributions were normalized by calculating the percentage frequencies for each 0.5°C interval represented. 408 Results Larval Survival Mortality in all groups was high (10%) throughout the first 10 days, until we reached a stable level of about 60-100 individuals per 36-L aquarium. With experimental requirements and additional mortality, the supply of larvae was exhausted 24 days after hatching. Thermal Selection The tendency to congregate within a particular temperature range was manifested in newly hatched larvae and did not vary significantly throughout development. This condition was maintained from 12h to 24 days after hatching (Table 1). The mean temperature selected fluctuated quite markedly, e.g., 3.1°C range for Group A, and 1.9 and 1.3°C for A, and B. There was no observed effect of acclimation temperature on temperature selected during the first 3 days after hatching. THE CANADIAN FIELD-NATURALIST Vol. 91 Behavioral Observations Until swim bladder inflation occurred between 7 and 10 days after hatching (Group A), 23°C), vertical position in the gradient and aquaria was regulated through a matching of upward swimming force (short bursts of rapid tail beating), and passive negative buoyancy (sinking). Larvae maintaining constant depth would therefore normally be slightly inclined with the head directed upwards (Figure la). To ascend, larvae increased their angle of inclination (Figure Ib) or accelerated locomotory movements. Both mechanisms were actually observed, separately and in combination. When activity ceased, the greater density relative to water, together with an anterior center of gravity, caused immediate descent with the head directed towards the bottom (Figure Ic). The rate of sinking was about 1.5 cm/s. The common behavior pattern appearing in all selection experiments prior to inflation of the swim bladder consisted of the following: (1) slow ascent into constant depth ascent descent FiGurE |. Orientations adopted by yellow perch larvae prior to swim bladder inflation. Vectors indicate negative buoyancy (dashed), active propulsion (light solid), and net force (heavy solid). OWE warmer water, (2) increased activity upon encounter- ing elevated temperatures, (3) very rapid penetration to the surface where the fish swam rapidly ina random path, apparently attached to the surface film, (4) sudden cessation of all activity, causing descent into cooler water, followed by (5) revival, resumption of activity, and re-ascent. After repeated ascents into very warm water (> 30°C), some larvae did not revive and drifted slowly to the bottom remaining there for the duration of the experiment. If the surface of the gradient was extremely warm (> 40°C) larvae would become immobilized before reaching the surface. An interesting side effect of this behavior occurred in larvae from Group A, (25°C) which had appar- ently injested air bubbles. The expansion of gas con- tained in the gut resulted in a strong positve buoyancy as the fish entered warm water, ultimately resulting in a situation in which the fish lay ventral-side-up on the surface, unable to regain depth. Coincident with normal inflation of the swim bladder the behavior described above was replaced by one which was more typically adult. That is, vertical position was regulated primarily by bladder buoy- ancy, swimming being reserved for locomotion. Constant depth could be maintained without activity and, as a consequence, the behavior of larvae in the gradient was less dynamic after swim bladder inflation than before. Fish remained longer in one place (at one temperature). Activity tended to decrease with age, but this observation was not quantitatively sub- stantiated. Discussion The most striking feature of our frequency distributions is that they were somewhat broad- shouldered (platykurtotic). The data of McCauley and Read (1973), although not strictly comparable, indicate that much more precise selection occurred in adults and juveniles; under-yearlings captured in October (5 months old) and acclimated to 24°C congregated with some precision (standard deviation —1.8°C) around 23.3°C. The mean standard deviation obtained here was 3.5°C (Table 1). This may be a result of a relative insensitivity of larvae to tempera- ture and may be related to the development of cut- aneous temperature receptors or of neural connec- tions, or both. The lack of any detectable acclimation effect during the first 3 days after hatching may be attributable to the small range of acclimation temperatures repre- sented. The slope of the preferred temperature- acclimation curve from 20° to 25°C for juveniles quoted by Ferguson (1958) was very small; for an acclimation range of 20° to 25°C the corresponding range of preferred temperatures was 23.1 to 24°C NOTES 409 (final preferendum = 24.2°C). These data are in keeping with the present observation, given the lack of precision with which larvae exhibited temperature preference. The behavior of individual larvae prior to swim bladder inflation must be considered maladaptive in situations where surface temperatures have been artificially elevated by thermal effluents. The tendency to penetrate very hot water and to become im- mobilized as a consequence, has also been observed in larval lake whitefish (Coregonus clupeaformis) by Hoagman (1974). The consistency of this behavior pattern together with its occurrence in at least one other species, suggests that it may have some utility in a natural situation, possibly in response to gentler gradients. The high temperatures encountered in the experimental situation described here may, through overstimulation, trigger some fixed-action pattern which causes larvae to ascend. The frenzy observed in association with this behavior is probably in part an’ expression of the effects which high temperatures have on biological functions in general, according to the principles of Arrhenius. Acknowledgments Thanks are due to Nancy McKerracher and Michael Prendergast, who helped with the field collections; and to Wayne King, who provided technical advice and assistance. We also thank Dan Faber and Fred Fry who gave helpful suggestions and, upon its completion, read through our manuscript. Literature Cited Everopeyzera, N. B. 1944. Preferred temperatures of fish larvae, Comptes Rendus de !’Academie des Science de l Academie de !’Union de la République Soviete Socialiste 42: 137-141. Ferguson, R. G. 1958. The preferred temperature of fish and their mid-summer distribution in temperate lakes and streams. Journal of the Fisheries Research Board of Canada 15: 607-624. Fry, F.E.J. 1937. The summer migration of the cisco, Leucichthys artedi (LeSueur), in Lake Nipissing. Uni- versity of Toronto Studies, Biological Series 44. (Publi- cations of the Ontario Fisheries Research Laboratory 55.) 91 pp. Hoagman, W. J. 1974. Vital activity parameters as related to the early life history of larval and post-larval lake whitefish (Coregonus clupeaformis). In Early life history of fishes. Edited by J.H.S. Blaxter. Springer-Verlag, Heidelberg. pp. 547-558. Hokanson, K.E.P. and C.F. Kleiner. 1974. Effects of constant and rising temperatures on survival and develop- mental rates of embryonic and larval yellow perch, Perca flavescens (Mitchill). Jn Early life history of fishes. Edited by J.H.S. Blaxter. Springer-Verlag, Heidelberg. pp. 437-448. Hoss, D. E., W. F. Hettler, Jr., and L. C. Coston. 1974. 410 THE CANADIAN FIELD-NATURALIST Effects of thermal shock on larval estuarine fish. Ecological implications with respect to entrainment in plant cooling systems. Jn Early life history of fishes. Edited by J. H.S. Blaxter. Springer-Verlag, Heidelberg. pp. 357-371. Houde, E. D. 1969. Sustained swimming ability of larvae of walleye (Stizostedion vitreum) and yellow perch (Perca flavescens). Journal of the Fisheries Research Board of Canada 26: 1647-1659. McCauley, R. W. and L. A. A. Read. 1973. Temperature Vol. 91 selection by juvenile and adult yellow perch (Perca flavescens) acclimated to 24°C. Journal of the Fisheries Research Board of Canada 30: 1253-1255. Powles, P.M. 1974. Survival of Australian Anchovy (Engraulis australis) eggs and larvae in a heat trap. In Early life history of fishes. Edited by J. H. S. Blaxter. Springer-Verlag, Heidelberg. pp. 373-381. Received 25 May 1977 Accepted 23 August 1977 Gestation Period and Juvenile Age at Emergence in Richardson’s Ground Squirrel GAIL R. MICHENER Zoology Department, University of Alberta, Edmonton, Alberta T6G 2E9 Michener, Gail R. 1977. Gestation period and juvenile age at emergence in Richardson’s Ground Squirel. Canadian Field- Naturalist 91(4): 410-413. Field observations of individually marked animals, and rearing of captive pregnant animals, provided information on time of breeding, length of the gestation period, and age of juveniles at emergence in Richardson’s Ground Squirrels. Female squirrels are mated within a couple of days of spring emergence from hibernation, young are born after a 24-day gestation period and first leave the natal burrow 30 days after birth. Key Words: breeding, ground squirrels, gestation period, hibernation, juvenile emergence, Sciuridae. Confusion exists in the literature as to the length of the gestation period in Richarson’s Ground Squirrel, Spermophilus richardsonii. Denniston (1957) esti- mated gestation to be 17 days, based on an observed copulation of a recently captured female and her subsequent parturition. Asdell (1964) stated gestation to be 28 to 32 days, based on Howell’s (1938) report of unpublished data on captive squirrels. Nellis (1969) and Sheppard (1972), quoting Asdell (1964) as their source, assumed a 28-day gestation when calculating conception dates of Richardson’s Ground Squirrel embryos. The variation in reported gestation periods suggests that further information is needed before these estimates are used as predictors of time of breeding or time of parturition. Because copulations are rarely seen in the field, and because laboratory breeding is not usually successful, estimates of the gestation period of Richardson’s Ground Squirrels are most readily obtained by indirect means. Data collected during studies on the effect of climatic conditions on spring emergence from hibernation and on breeding (Michener 1973, 1977) indicated that the gestation period falls between the 17-day and 28-day estimates. This note elaborates on the data I previously collected, and presents additional data on the length of the gestation period, as well as information on the time of breeding with respect to female emergence from hibernation, and the age of juveniles at emergence from the natal burrow. Eight, 31, 23, and 16 female Richardson’s Ground Squirrels were live-trapped in mid-April of 1969, 1970, 1971, and 1972, respectively, in southern Saskatch- ewan. Most of these squirrels (60 of 78) were collected within a 320-ha prairie pasture near Kayville, Saskatchewan (49°40’N, 105°10’W, elevation 780 m). The squirrels were individually caged. Details on caging and diet can be found in Michener (1971). Each animal was checked daily and litters were recorded as being born on the day on which they were first observed. Sixty-five of the females produced litters; of these 38 had been captive for 17 or fewer days and 27 for more than 17 days (Table 1). Thus 42% of the females had been held captive, separated from males, for longer than the 17-day estimate of the gestation period made by Denniston (1957). Since Denniston observed copulation between his captive female and a male, and since the weights reported forthe young of this female indicate that the litter was not premature, the female was presumably pregnant at capture and the mating was aberrant behavior associated with the stress of captivity. Two of my squirrels gave birth after 23 days in captivity (Table 1), and this period is taken as a minimum estimate of the gestation period. These data do not allow a maximum estimate of the Sy NOTES 411 TABLE |—Frequency distribution of the number of days in captivity prior to parturition, and of the number of days between earliest possible spring emergence and parturition Number of days / Y va . y iy Lower Arrow GF SS SS Z \ \ 7 . Lake \ \ 7 LEGEND ~ i \ y) a Copture Location il wh if | Release Location (0) we Sle ——————— UG (Champion Lakes o) Gp) ) \—__ Park co Y KS ) : ‘i \ FIGURE 3. Movements of Bear Number 478. May is limited, forage at a sufficient distance from low-elevation populated areas is scarce, and the transport of bears to high elevations is blocked by snow or washed-out roads. Recaptures of bears released in October were normally made the following year. Experience indicates that cubs left behind and good feeding areas act as the strongest incentives for bears to return, although this is difficult to demon- strate from the data. Bears were found to have travelled distances ranging from 10 to 99 map-km (straight-line distance), at rates of up to 11 km per day. In some instances, the apparent rate of travel was much slower. The topography was generally mountainous and rugged and this may have influenced the rate and direction of return for certain individuals. Bears were found to have crossed bodies of water approximately | km wide, but observations of three tagged bears indicate a tendency for the animals to walk around Kootenay Lake, which measures approximately 5 km across. Thirty-seven bears out of 54 were recaptured at their original capture location, nine of them making two or more returns from a variety of release sites. Distances travelled and rates of return varied greatly. Some bears were recaptured at the original capture location as long as one year after release and may have 422 THE CANADIAN FIELD-NATURALIST returned to the capture site at once. Ten bears, however, made their returns to original capture locations within one month of release. Bear Number 228, a female with cubs, was originally captured on7 July 1971 at Champion Lake Park (Figure 1). She and one cub were moved; the second cub was destroyed. From a release location 38 km away at Clearwater Creek, this adult and her cub returned to Champion Lake Park. They were recaptured on 15 July 1971 and released 88 km away at Fletcher Creek. On 3 June 1972 this female was recaptured for the third time in the park and released 99 km away at Retallack. On 11 June 1972 she was back at the park for a fourth time. This time she was released across Kootenay Lake at Crawford Creek, 88 km away. On I5 June 1972 she was shot at Arrow Creek and it is possible that she was in the process of returning once more. Bear number 19 returned to the original capture location (a slaughter pit) following two different releases. The first return was a trip of approximately 22 air-km, the second, more than 56 km, was accom- plished in 7 days (See Figure 2). Bear number 478 illustrates that the return to Vol. 91 original capture location is not necessarily simply a return to the nearest source of garbage. This bear was released within 8 km of the garbage dump at Nelson, but was recaptured at its original capture site more than. 32 km from Nelson (See Figure 3). The data indicate that dump use is not dependent on simple random movement. If we consider bear mobility and the extensive capability of bears to return home, open garbage dumps pose a serious game management problem. Acknowledgments We are most grateful to Gordon Hartman for his encouragement and advice during preparation and finalization of the paper, and to Anne I. Dagg for her assistance with the early stages of the original manuscript. We appreciate the work of Timothy Rutherglen, who provided information and assistance during 1972 and Peni Campbell, who worked on the figures. Dubious thanks should go to the communities of the West Kootenay that have provided garbage dumps and the garbage bear problem. Received 14 August 1974 Accepted 19 August 1977 Paspalum ciliatifolium, a Grass New to Canada from Southwestern Ontario WILLIAM J. CRINS,! PAUL D. PRATT,?2 and DANIEL F. BRUNTON} 11336 Bunnell Drive, Burlington, Ontario L7P 2El 2Box 69, Benheim, Ontario NOP 1A0 3Box 757, Canmore, Alberta TOL 0MO Crins, W. J., P. D. Pratt, and D. F. Brunton. 1977. Paspalum ciliatifolium, a grass new to Canada from southwestern Ontario. Canadian Field-Naturalist 91(4): 422-424 The grass, Paspalum ciliatifolium Michx., is reported as a new addition to the flora of Ontario and Canada. The original discovery was made in Essex County in 1975, with two additional stations being found in Kent County in 1976. The ecology and taxonomy of the species are discussed and it is suggested that P. ciliatifolium will likely be found elsewhere in southwestern Ontario. Key Words: Paspalum ciliatifolium, new to Canada, Ontario, distribution, grass. On 31 August 1975, the authors discovered a large station of the grass, Paspalum ciliatifolium Michx., at Windsor, Essex County, Ontario. Subsequently, on 18 September 1976, in adjacent Kent County, P.M. Catling, A. A. Reznicek, R. Brown, and S. M. McKay located two additional stations of the species (see Figure |). The genus Paspalum is not listed by Soper (1949) nor by Boivin (1967), and examinations of various herbaria (including CAN, DAO, OAC, and TRT) have turned up no records. As we could find no evidence of its previous occurrence in the country, we conclude that the Essex County station constitutes the first record of any member of the genus, not only within Ontario, but from anywhere in Canada. Taxonomy The species here referred to as P. ciliatifolium Michx. has been the subject of differing taxonomic interpretation. In view of the fact that the material has been identified at the varietal level, a brief discussion 1977 FIGURE 1. Distribution of Paspalum ciliatifolium var. muhlenbergii (@) and Paspalum ciliatifolium var. stramineum aD in Ontario. of these taxa is appropriate. Several early works dealing with the genus Paspalum have considered P. ciliatifolium to consist of three separate species (Nash 1912; Rydberg 1932; Hitchcock and Chase 1950). Gleason (1952), however, recognizes a single species, P. ciliatifolium, composed of three varieties cor- responding to the three species of earlier authors. It is Gleason’s interpretation that we are following. Voss (1972) indicates the presence of two varieties of P. ciliatifolium in Michigan, P. ciliatifolium var. muhlenbergii (Nash) Fern. and P. ciliatifolium var. stramineum (Nash) Fern. Both of these varieties have been found in the Ontario collections described here. Species Range In the broad sense, P. ciliatifolium is a widely distributed species of the United States (Hitchcock and Chase 1950; Gleason 1952). The American range is described by Gleason (1952) as extending from New Hampshire and Massachusetts, west to Michigan, Minnesota, Kansas, and Arizona, and south to the Gulf of Mexico. Paspalum ciliatifolium var. muhlen- bergii is found throughout much of this range, but P. ciliatifolium var. stramineum tends to occur in the western part of the range, being considered a species of the prairies and plains (Rydberg 1932). The first Canadian station, in Essex County, was revisited on 16 August 1976, when voucher specimens were collected (deposited in OAC, TRT). Specimens of the Kent County stations have been deposited in TRT. Collection data for the Ontario stations are as follows: Z P. ciliatifolium var. muhlenbergii Essex Co.: East end of Rickard Street ca. 320 m west NOTES 423 power line, west of Malden Road, Windsor (in Ojib- way Prairie Provincial Nature Reserve); open, dry, clay soil beside overgrown roadbed, forming dominant ground cover. W.J. Crins, P. D. Pratt, H.L. Dickson, J. Goltz, and R.J. Pittaway 761. 16 August 1976 (OAC, TRT). Kent Co.: Zone Township, ca. 1.6 km east of Bothwell, 42°39’N, 81°51’W; open, sandy soil along roadside with Festuca rubra. P. M. Catling, A. A. Reznicek, R. Brown, and S. M. McKay. 18 September 1976 (TRT). P. ciliatifolium var. stramineum Kent Co.: ca. 3.2 km north of Thamesville Station, west side of County Road 26 (to Florence), 42°37'N, 82°0.5’W; dry, sandy soil in open meadow. P. M. Catling, A. A. Reznicek, S. M. McKay, and R. Brown. 18 September 1976 (TRT). Habitat At the Essex County site, the species forms a dense ground cover over disturbed clay soil beside a little- used road. It was found in open areas as well as under the partial shade of Pin Oak (Quercus palustris Muench.) on an old, unused track nearby. In 1975 and again in 1976, the species was observed in other sites within the Ojibway Prairie Provincial Nature Reserve (known also as the “Windsor Prairie’). The three Ontario stations appear to be quite similar. All are on dry, disturbed, relatively open sites. These areas have long been known for their relict prairie flora (see Rogers 1966), so P. ciliatifolium is a consistent addition to the flora of the area. In view of the relatively large extent of this type of habitat in southwestern Ontario, we expect that P. ciliatifolium will be found in other localities in that part of the province. Acknowledgments The authors express their appreciation to P. M. Catling and A.A. Reznicek of the University of Toronto for supplying valuable comments, data, and verification of the variety of the Essex County specimen. We also appreciate the opportunity of searching the CAN, DAO, OAC, and TRT herbaria, and thank their curators, J. M. Gillett, W. J. Cody, J. F. Alex, and P. M. Catling, respectively. Literature Cited Boivin, B. 1967. Enumération des plantes du Canada. VI. Monopsides. (Deuxieme partie.) Naturaliste Canadien 94: 471-528. Gleason, H. A. 1952. The new Britton and Brown illus- trated flora of the northeastern United States and adjacent Canada. 3 volumes. New York Botanical Garden, New York. Hitchcock, A. S.and A. Chase. 1950. Manual of the grasses of the United States. 2nd edition. United States Depart- 424 ment of Agriculture Miscellaneous Publication 200. 1051 pp. Nash, G.V. 1912. Paspalum. North American Flora 17(2): 165-196. Rogers, C. M. 1966. A wet prairie community at Windsor, Ontario. Canadian Field-Naturalist 80: 195-199. Rydberg, P. A. 1932. Flora of the prairies and plains of central North America. New York Botanical Garden, New York. THE CANADIAN FIELD-NATURALIST Vol. 91 Soper, J. H. 1949. The vascular plants cf southern Ontario. Federation of Ontario Naturalists, Toronto, Ontario. 95 pp. Voss, E.G. 1972. Michigan Flora. Part 1: Gymnosperms and monocots. Cranbrook Institute of Science Bulletin 55. 588 pp. Received 31 March 1977 Accepted 20 September 1977 Additional Record of the Southern Flying Squirrel from Quebec D. J. OXLEY and J. M. GALL Vanier College CEGEP, Snowdon Campus, 5160 Decarie Blvd., Montreal, Quebec H3X 2H9 Oxley, D.J. and J.M. Gall. 1978. Additional record of the Southern Flying Squirrel from Quebec. Canadian Field-Naturalist 91(4): 424. During a study of small mammal movements at Pine Hill, Argenteuil County (45°44’N, 74°29’W), Quebec, four Southern Flying Squirrels, Glaucomys volans, were captured in Sherman traps on a 0.9-ha sample plot between 19 and 26 October 1976. The trap-site was a well drained, mature beech-maple (Acer-Fagus) forest, which supports the highest populations of G. volans in Canada (Banfield 1974). Previous records from Quebec include one specimen collected south of the Ottawa River at Hudson, Vaudreuil County, and four specimens from Gatineau County northwest of Hull (Youngman and Gill 1968). Three of the squirrels were marked and set free. The fourth was retained as a specimen in the Vanier College Museum of Natural Science (VCMNS - 301). External measurements in millimetres were 224, 91, 29, 20; weight was 55 g and the testes were abdominal. abdominal. Literature Cited Banfield, A. W. F. 1974. The mammals of Canada. Univer- ‘sity of Toronto Press, Toronto. xxv+ 438 pp. Youngman, P. M. and D. A. Gill. 1968. First record of the southern flying squirrel, Glauwcomys volans volans, from Quebec. Canadian Field-Naturalist 82: 227-228. Received 22 March 1977 Accepted 19 September 1977 Occurrence of the Green Sunfish (Lepomis cyanellus) in the Grand River System EDWARD KOoTT and GREGORY B. HUMPHREYS Department of Biology, Wilfrid Laurier University, Waterloo, Ontario N2L 3C5 Kott, Edward and Gregory B. Humphreys. 1977. Occurrence of the Green Sunfish (Lepomis cyanellus) in the Grand River system. Canadian Field-Naturalist 91(4): 424-426. The Green Sunfish (Lepomis cyanellus), a species of restricted distribution in Ontario, is reported from the Nith River of the Grand River drainage basin. This species has previously been recorded only from the Thames River drainage system of southwestern Ontario, and in some lakes in Quetico Park in northwestern Ontario. The possibility of species transfer from the Thames system to the Grand system is discussed. The Green Sunfish (Lepomis cyanellus) has a restricted distribution in Ontario. It has previously been found only in several lakes of Quetico Park in northwestern Ontario and in the Thames-Avon River system of southwestern Ontario (Scott and Crossman 1973; Crossman 1976). The present note records the occurrence of this species from a second drainage basin in southwestern Ontario. During a seining program in the Nith River, a tributary of the Grand River, several specimens of L. 1977, NOTES 425 FiGure |. Lateral view of the Green Sunfish (Lepomis cyanellus), 157 mm total length (128 mm standard length), collected from the Nith River on 10 June 1976. cyanellus were collected at two locations. One locality was at a washed-out dam just north of Plattsville in Oxford County. On 10 June 1976, eight specimens were collected; on 30 September 1976, 14 individuals were collected. Both collections were from a deep (over 1.9m maximum depth) oxbow with only a limited connection with the river at the time of collecting. The bottom was muddy. Specimens from the 10 June collection ranged in size from 44.5 mmto 157 mm total length (37.2 mm to 128 mm standard length). Specimens from the Sep- tember collection ranged in total length from 31.2 mm to 90 mm (26 mm to 73 mm standard length). The second location where specimens were col- lected was at Mornington Centennial Park, just north of Millbank, Perth County, on 25 March 1976. Four fingerlings were caught. The specimens were collected over a flood plain during a period of high water. The fingerlings ranged in size from 32.9 mm to 37.4 mm total length (25.4 mm to 29 mm standard length). Adult L. cyanellus can be separated from other species of Lepomis by the large mouth, large eyes, and relatively short rounded pectoral fins (Figure 1). A dark spot is usually present at the base of the last rays of the dorsal fin, a character it shares with L. macrochirus, the bluegill. Specimens up to about 30 mm in total length, when alive, have a lateral coloration consisting of vertical rows of darkened spots enclosed within a vertically elongated light blue halo. This color pattern largely disappears when a specimen is placed in preservative. Slightly larger specimens do not exhibit the banded coloration of smaller individuals. No other specimen of Lepomis was collected in the Nith River. Lepomis gibbosus, the Pumpkinseed, however, has been recorded from Horner’s Creek (= Whiteman’s Creek, Mayall 1954), which is the first major tributary of the Grand River south of the Nith River. It flows in a general southeasterly direction as does the Nith. More extensive sampling may reveal the presence of the Pumpkinseed in the Nith River. The only other centrarchids collected in the Nith River were the Rock Bass (Amb/oplites rupretis) and the Smallmouth Bass (Micropterus dolomieut). The Green Sunfish has been reported from the Thames-Avon River system, which drains into Lake St. Clair. An examination of the topographic map Stratford 40P/7, edition 5, 1972, published by the Department of Energy, Mines and Resources, Ot- tawa, shows that Silver Creek, a tributary of the Nith River, is separated from a series of ditches that drain into the Avon River by a distance of 160 m (at 43°24’25” N and 80°51’12” W). On 14 March 1977, a period of high water, the area of separation between the systems was visited. At this time the two systems were connected by a ditch in which the water was about 1.9 m wide and 23 cm deep and the flow was towards the Nith River. Following the ditch south- westward towards the Avon River, the flow in the ditch was still northward towards the Nith River through an extensive low-lying flooded area. This ditch eventually connected with a second ditch whose water flow was southward into the Avon River. At high water, therefore, at least one connection exists between the Avon River and the Nith River. This provides a route for the transfer of the Green Sunfish from the Thames-Avon system to the Grand-Nith system which drains into Lake Erie, and suggests a possible reason for its occurrence in the Nith River. The specimens of L. cyanellus are catalogued in the Wilfrid Laurier Museum as collections WLU 4557, 426 4696. and 4911 and in the National Museum of Natural Sciences as NMC 77-0610. We thank Frank Mallory for his help in collecting the specimens. The specimens were verified as L. cyanellus by G. C. Gruchy of the National Museum of Natural Sciences, Ottawa. Literature Cited Crossman, E. J. 1976. Quetico fishes. Royal Ontario Mu- THE CANADIAN FIELD-NATURALIST Vol. 91 seum, Life Sciences Miscellaneous Publications. ix + 86 Pp. Mayall, K. M. 1954. Part I] — Wildlife. Grand River Conservation Report. 1954. Ontario Department of Plan- ning and Development, Conservation Branch, Toronto. Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada, Bulletin 184. xi + 966 pp. Received 23 March 1977 Accepted 23 July 1977 Book Reviews ZOOLOGY The Red Colobus Monkey By Thomas T. Struhsaker. 1975. Wildlife Behavior and Ecology Series, University of Chicago Press, Chicago. 311 pp., illus. $25. In collecting the material for this book, the American primatologist Thomas Struhsaker spent 1593.7 hours observing Red colobus Monkeys in the Kibale Forest Reserve of western Uganda, writing his field notes using carbon paper so that there was little possibility of their loss. He solved the difficulty of working in a dense forest by observing the monkeys from trails marked on the ground ina grid pattern cut along compass bearings, whose intersections occurred either at 50- or 100-m intervals. His book incorporates many of his raw data, making it a more detailed work than the earlier ones in the same series, such as The Serengeti Lion by George Schaller and The Spotted Hyena by Hans Kruuk, and therefore one less appealing to the layman. In addition to the descriptive text some of the data are incorporated into 58 detailed tables. Red Colobus Monkeys weigh about 10 kg, with the males somewhat heavier than the females. They are not really red, but have large or small patches of chestnut-colored fur on their trunks, depending on the subspecies. Like most monkeys the Red Colobus lives in social groups that include more females than males. Although they are considered to be arboreal animals, they tend to move rather clumsily in the trees, often jumping from one tree or branch to another with a mighty spread-eagled leap. An interesting feature of this book is the extensive treatment of vocalizations. Struhsaker used a tape Reptiles and Amphibians in the Service of Man By Wilfred T. Neill. 1974. Pegasus, New York. 248 pp. The sponsors of this book’s series (Science and Society) promise at the outset that this biological sciences curriculum study book will be “highly readable,” “nontechnical,” and instructive. It 1s. Neill, noted Florida-based herpetologist and author of the superb Last of the Ruling Reptiles, has produced another fine book. Even the title is provoca- tive — surely we do not usually think of reptiles and amphibians as serving man. In this book, Neill is lucid, persuasive, interesting, and often speculative. He frequently argues by anecdote and analogy, and he argues from wide field and laboratory research (see recorder to tape 16 discrete call types, many of them illustrated by sonagrams. They are described in a number of sections with intriguing titles such as Scream, Sqwack, and Shriek; Wheet; Bark-chist and Chist-bark; Uh!; and Squeals and Gasp of Dying Infant (the infant was found lying neglected on the ground, so Struhsaker carried it home and before it died made extensive recordings of its cries). In this 40- page chapter on vocalizations, as in the others, Struhsaker employs a comparative style that greatly increases the value of his work. After describing the basic sounds made by Colobus badius tephrosceles monkeys of Uganda and noting when, by whom, and under what circumstances they were made, he compares their repertoire with that of other races, and his analyses with those of other primate workers. Zoologists studying other species will find these summaries invaluable. Struhsaker hopes with the publication of this work that more biologists will become interested in the rain forest biome, not only because it is a fascinating, little- studied habitat, but because it is being rapidly destroyed in tropical countries throughout the world. Since it would take 200 or 300 years to replace the largest trees, this biome, once lost, will probably never be regained. Rain forests supply some lumber, but their value as a source of oxygen and of tourism is far greater. ANNE INNIS DAGG Box 747, Waterloo, Ontario N2J 4C2 p. 85). In fact the twenty chapters are laced with personal observations. More important, and for the sake of brevity, he does not just retell the sorry tales of over-exploitation of so many “herptiles” by man (“herps” Neill considers slang, p. 5). Periodically, he debunks myths about his subjects, or reveals that facts about them are often stranger than fiction (e.g., p. 146). Technical “hardware” and technology are usually well incorporated at appropriate points: in the treatment of developmental biology of the frog and the space program, radio-telemetry, computerized re- trieval systems, etc. His sense of herpetofauna and ecological associations is keen, especially when he 427 428 discusses such little-studied topics as the effect of controlled burning (chapters III-IV). His writing also has a highly graphic quality, rather like that of Carl Kauffeld and Raymond Ditmars, which creates con- siderable immediacy. Also, the quite plentiful black- and-white photographs, often simple, may be dramatic (see Figures 29 and 30 illustrating the effects of snake bite). Some behavioral theories are briefly but intrigu- ingly examined, such as the impulse of hatchling turtles “to fight gravity” (p. 171), and pond specificity (p. 188) of certain amphibians. To a point, an inter- disciplinary approach is used, when dealing with cancer and radiation (p. 166), and discoveries re- garding sleep (pp. 214-215). Neill suggests directions for herpetological research to take (pp. 75, 168) and problems in such research (p. 117). The Canadian reader will glean interesting data. For example, on p. 159 we learn that the burrowing “common hog-nosed snake” (no scientific name given) has little or no melanin in its peritoneum, and A Field Guide to the Birds of West Africa By W. Serle and G. J. Morel. Illustrated by W. Hartwig. 1977. Collins, London. 351 pp. £5.95. This latest field guide to birds published by Collins is a case of good intention gone awry. For years, ornithologists and bird-watchers travelling to West Africa have needed a pocketable book to bridge the gap between Bannerman’s The Birds of Tropical West Africa and Elgood’s Birds of West African Town and Garden. A Field Guide to the Birds of West Africa ought to have filled this gap, but the book has some serious shortcomings which limit its utility. At first sight Birds of West Africa appears to comply with the accepted, modern-day standards set for field guides. It is concise, clearly printed, full of what seems to be informative text, and is illustrated with 48 plates, 28 of them in color. There is a useful introduction to the topography, climate, and vegeta- tion of West Africa at the beginning of the book, and two checklists at the back. One is a checklist of species, the other, a list of scientific and vernacular names in English, Spanish, French, and German. Species in the text are treated under the headings of Identification; Voice, Habitat and Nesting. Notes on allied species are given where applicable. A total of 726 species is given treatment (or mention), and of these, 515 are illustrated, 335 of them in color. The remaining 371 of 1097 West African species are included in the checklist already mentioned. In using the book, one finds the errors and annoying inconsistencies, which for a field guide, can hardly be tolerated. Consider the _ illustrations. THE CANADIAN FIELD-NATURALIST Vol. 91 on p. 180 that queen snakes are not very difficult to capture. The book is full of facts without being cluttered. In some places, however, a little more detail would be welcome, as for example, regarding the recent discoveries of new species and races of reptiles and amphibians within the United States (pp. 9-11). Challenging questions are often posed, but sometimes left unanswered. Perhaps on occasion, Neill is too speculative, or too general. More on reptiles and amphibians as indicators of environmental quality and on adaptations to urbanization would be helpful. But specifically, perhaps the worst that can be said of this excellent book is that the style is sometimes abrupt (chapters VIII to IX, “from ecology to enzymes”), and the author too philosophical, as for example when he appears to be anti-women’s lib (p. 235). C. A. CAMPBELL 421 King Street, Waterloo, Ontario N2J 3Z4 Preference is often given to the illustration of species found in one of the authors’ own territories (Senegal for Morel, Cameroun for Serle). Thus we find that the Striped Swallow depicted on plate 35 is of the race maxima, restricted to Cameroun, whereas the more common and widespread form, puella, different in appearance, is not shown. Similarly, the male White- crowned Cliff-chat in Senegal lacks the white crown, but in the remainder of its range, the male possesses it, and we find the dark-crowned form illustrated. One wonders too, when space for illustration is at a premium, why a species such as the House Martin, an uncommon palaearctic winter visitor to West Africa, is illustrated at the expense of common African species such as the Red-rumped and Rufous-chested Swallows. Most plates show an alarming disregard for scale. The Brubru Shrike on plate 36 is shown to be marginally smaller than the Long-tailed Shrike, yet the former is five-and-a-half inches in length, and the latter, twelve. Such errors of scale can be found on virtually every plate. The proportions of some birds are not correct: the tails of bee-eaters and the heads of owls, to mention only two. The guide also fails I think, to deal adequately with the difficult groups of West birds, notably cisticolas and bulbuls. Many cisticolas haye distinct breeding and non-breeding plumages, and these are given full treatment in the text, but the black-and-white plate devoted to this group of birds is unsatisfactory. In the case of bulbuls, I feel the text falls short. The forest pycnonotids are notoriously 1977 difficult to observe, let alone identify. Song is the surest clue in most cases. Yet for two “troublesome” species we are expected to distinguish “bursts of excited throaty babbling” from “bursts of concerted throaty babbling.” Some oddities in nomenclature crop up from time to time, perhaps the most obvious one being the naming of Pogoniulus chrysoconus as the Yellow- fronted ‘Barbet,’ when small capitonids of the genus Pogoniulus are generally referred to as ‘tinker-birds.’ All other pogoniuline barbets in the text are called tinkerbirds. There is a case of text not matching illustration. A good field mark for the Rock Thrush is the white lower back and rump, and this is properly pointed out Vertebrate Biology By Robert T. Orr. 1976. Fourth Edition. W. B. Saunders, Philadelphia. 472 pp., illus. $13.35. That this is a superior text on vertebrate biology goes without saying; any book that is so popular that it has gone into three new editions since it was first published in 1961 and which has been translated into Spanish must be fulfilling a need for a basic zoology text. In each succeeding edition, the author, Dr. Orr of the California Academy of Sciences, has updated the text, to a limited extent in the sections on anatomy, and to a large degree in the discussions of ecology, ethology, and population dynamics, fields in which new information is being discovered more rapidly. The first half of the text considers the major classes of vertebrates; the second deals with general topics such as systematics, distribution, reproduction, dormancy, and growth and development. Most, but certainly not all, of the animals discussed live in North BOTANY BOOK REVIEWS 429 in the text, but not shown in the illustration. Which brings up my final point. In a book credited in the preface to follow the Peterson system of identifica- tion, there is not a single use of the very useful field- mark pointers on the plates. It is a pity that in this era of field guide excellence there appears one which falls short of the mark. But for people in West Africa who wish to venture beyond Elgood’s Birds of West African Town and Garden, this new guide is just the job. MICHAEL DYER Department of Zoology, University of Aberdeen, Aberdeen, Scotland AB9 2TN America, and there are a number of sections which will be of special interest to Canadians. These largely concern our arctic fauna, dealing with phenomena such as migration, hibernation, and cyclic fluctua- tions of some vertebrate populations. Historically Orr considers the effect the Ice Age and attendant glaciation had on the distribution of animals in Canada and on their systematics. The many black-and-white photographs that illustrate the subject matter of this book are excellent, as are the figures depicting various specific features of animals. Indeed the entire book is a fine one, well worth considering as a text by any professor who teaches a course in vertebrate biology, or by any biologist who needs a reference book on vertebrates. ANNE INNIS DAGG Box 747, Waterloo, Ontario N2J 4C2 Seeds and Fruits of Plants of Eastern Canada and Northeastern United States By F. H. Montgomery. 1977. University of Toronto Press, Toronto. 232 pp., illus. $25.00. Professor Montgomery has done it again! He has produced another invaluable aid to the identification of our flora, which is well keyed and extensively illustrated. Until the appearance of this book bot- anists and others who wanted to identify a seed, and did not have the rest of the plant, had nowhere to turn and were dependent on the good offices of knowl- edgeable colleagues, or they had the tedious task of comparing the specimens to be identified with ones in a herbarium or seed collection. Professor Mont- gomery’s book will simplify the process and make seed and fruit identification available to all who have a modicum of botanical expertise. The book employs a novel and apparently original key based on gross morphology, and shape in longi- tudinal and cross sections. The seeds (or the fruits if these are indehiscent and the means of dispersal of the plant) are, for each of the species, keyed, described, 430 and illustrated. Eleven hundred species, both native and introduced, belonging to 118 families, are included. The illustrations are all black-and-white photographs of the whole seed or fruit, usually shown in several views. They vary considerably in quality but most are at least adequate for their purpose. It is unfortunate that details of surface structure are not shown, for only whole seeds or fruits are illustrated. The use of scanning electron microscope photographs to illustrate these details would have been most useful, for in many species they are diagnostic. Diagrammatic illustrations to show the precise meaning of the major botanical terms used in describing the seeds and fruits removes any possibility of ambiguity, and a full glossary covering almost all the terms used in the account is included. A good index to both English and Latin names enables the user to locate information Carnivorous Plants By Randall Schwartz. 1974. Praeger (Canadian distribu- tor Burns and MacEachern, Don Mills). 128 pp., illus. $8. Here is another popular book on carnivorous plants or, as many people will know them, insectiv- orous plants. This book is for the amateur, simply written and simply presented with 44 black-and-white photographs of a selection of the more widely known species. The full-page photographs are mostly close- ups of leaves or plants and do help to fill space and make for a more impressive book. One wonders whether it all couldn’t have been presented much more attactively in bulletin form. Carnivorous plants are a fascinating group of plants, often grossly misunderstood, possibly the reason the author has included a seven-page section on science fiction and mythology as they relate to this group of plants. There is also a five-page section of basic information on “How to grow Carnivorous Plants.” These plants are not easy to grow in the modern home or apartment. There is, however, an ENVIRONMENT THE CANADIAN FIELD-NATURALIST Vol. 91 rapidly. The book is well produced on a stout semi- gloss paper, and bound so that the pages lie flat wherever the book is opened. It is a business-like production well suited for use on the laboratory bench, with the illustrations appearing alongside the description of each species. The arrangement is taxonomic, permitting ready reference and the com- parison of related species. This is a book that will be indispensible to all whose work or interests require the identification of seeds and fruits of plants from eastern North America. J. K. MORTON Department of Biology, University of Waterloo, Waterloo, Ontario N2L 3G1 increasing interest in terrariums, which make for better habitats for these plants that require exacting control of temperature, humidity, and moisture levels. The photographs are mostly very mundane and unexciting, usually close-ups that do not include a scale. Neither do they allow one to gather any idea of the overall appearance of the plants. As this is an exciting group of plants, and a real challenge to amateur horticulturists, it is unfortunate that the author and his publishers could not have found a more exciting way to illustrate them. A list of sources is included, of which the author is one. Perhaps we should consider this his catalogue, albeit an expensive one. There are no Canadian sources listed although the Montreal Botanical Gordens is included in a list of places where carniv- orous plants can be seen. L. C. SHERK Sheridan Nurseries Limited, Etobicoke, Ontario M9C IAI Physiological Limnology — An approach to the physiology of lake ecosystems By H. L. Golterman. 1975. Elsevier, New York. xiv + 490 pp., illus. $51.95. Lately there has appeared a number of textbooks, G. A. Cole’s Textbook of Limnology and R. G. Wetzel’s Limnology to name only two, each seeking to update the subject originally described in the classic works by Ruttner and Hutchinson. Golterman has chosen to examine the physico-chemical characteris- tics of aquatic ecosystems and their regulating influences on the biochemistry and physiology of aquatic organisms. Although I did enjoy reading the book, and some 1977 chapters, such as Chapter 4 on primary production (photosynthesis) in relation to light under natural conditions, will certainly be appreciated by students and aquatic scientists who wish to familiarize them- selves with this complex subject, I could not help but feel that the book is already dated even considering its publication date. For example, Chapter 4 does not include the use of liquid scintillation techniques in !4C primary productivity studies. Liquid scintillation spectrometers are more efficient and convenient to use than the counting systems mentioned. No indication is made of the contribution to primary production of lakes by aquatic macrophytes although on page 2 It is noted that the algae are quantitatively the most important of the primary producers (macrophytes, algae, and photosynthetic bacteria), except within the littoral zone of lakes. Chapter 16 looks at bacterial limnology and the work by Wright and Hobbie on microbial utilization of dissolved organic compounds. No reference is made to the studies by Vaccaro, Jannasch and others who used Wright and Hobbie’s technique and found that it did not work under some conditions, e.g., usually in oligotrophic waters. Golterman notes that Wright and Hobbie made no correction for the loss of !4CO, by respiration in their experiments, but he does not cite the 1969 work of Hobbie and Crawford which does. They describe a method to measure the '*CO, produced and note the importance of it to the kinetic parameters measured in earlier work. Following the general introduction of Chapter 1, Chapters 2 and 3 cover lakes and their origins and the chemical composition of lakes. This is followed by a series of related chapters on the biological, biochemi- cal, and physico-chemical aspects of aquatic eco- systems. Not until Chapter 18 are lake sediments examined, almost as anafter thought. The importance of sediments to the physiology of lakes is appreciated by limnologists. I do feel that it would better have Mankind’s Future in the Pacific Edited by R. F. Scagel. 1977. University of British Colum- bia Press, Vancouver. The plenary and special lectures of the 13th Pacific Science Conference, August 1975. ix + 206 pp. Paper $6.95. Dr. Scagel has organized an impressive collection of 13 plenary and special lectures presented at the 13th Pacific Science Congress in August 1975. This inter- disciplinary collection, written for a general audience interested in catching up on many important issues now facing man in the Pacific region, represents the work of some of the world’s most prolific thinkers. The first two essays, written by Gerard Piel and Nathan Keyfitz, focus on the population problems in BooK REVIEWS 431 followed Chapter 3 so that in the following chapters references could have been made back to it. As it stands, the importance of sediments to the physico- chemistry and biology of the overlying waters would not be conveyed to students. This is not to say that the chapter is not interesting; indeed I found the chapter contained a great deal of interesting information. One last major criticism remains. In the preface the author notes that previous textbooks have not dealt with the questions that face the water manager, a valid point with which I heartily concur. Golterman then covers water management problems in the last chapter in some 20 pages. Hardly the last word on water management. I would like to have seen this chapter expanded considerably. In addition to Chapter 4, several other chapters were exceptionally well presented. These include Chapter 13 onalgaeand their pigments, Chapter 15 on energy and mass transport through food chains, and Chapter 17 on nutrient budgets and eutrophication. Chapter 13 is a good summary of the extensive literature to be found on algal physiology and both Chapters 15 and 17 are concluded with case studies that illustrate the theory discussed. Golterman intended, as outlined in the preface, that this book should be a first approach to relating the chemical environment with the biochemistry and physiology of the aquatic organisms in a quantitative way. Generally I believe he has accomplished this and that the book will be appreciated by the students for whom it was intended. Indeed, there is much in this book that all aquatic scientists could find useful and informative. It must be stressed that this book is not intended for lay naturalists but for advanced students and scientists. RICHARD D. ROBARTS 53 Selwyn Place, Kanata, Ontario K2K IPI the Pacific. Piel, who also writes about energy, quotes colleagues like Garrett Hardin, Paul Ehrlich, and Jacob Bronowski to substantiate his analysis of population growth. He does not, however, make any outstanding judgments or propose any solutions to the problems at hand. The essay by Keyfitz is a well- rounded analysis of the complicatcd variables that interact to shape today’s Pacific. In the next essay, geneticist L. H. Shebeski uses global food equations in a discussion about available food resources in the region. He confines most of his comments to land-based foods. M. Behar, a nutrition- ist, follows with an examination of the interrela- 432 THE CANADIAN FIELD-NATURALIST tionships of the food and poverty cycles. Energy, perhaps one of the most controversial topics of the last decade, is reviewed by Lord Ritchie Calder. In his essay, “All Life is Energy,” Lord Ritchie examines the sources and alternatives available to society. Technology, the mechanical extension of man’s brain, is reviewed by Maurice Strong, who writes . .. “Technology in the hands of industrial man can either enhance or destroy the natural environment resource base; it can either support or undermine the quality of human life in the region.” His theme is a very persuasive one in which the need for man to determine his own evolution is stressed. Further, he believes that “the struggle to preserve and enhance the environment of our ‘Only One Earth’ may be won or lost in the Pacific.” This might well be the case in view of the fact that the Pacific region is the largest in the world. Energy and technology are considered in many of the essays intermittently throughout the book. The social issues confronting man in the region are analyzed by Herman Kahn, a futurologist, and William Epstein, a specialist in arms control and disarmament. Epstein believes that weapons are a form of pollution to the human environment and unless man can ensure his own survival, then there is no point in attempting to improve the ecology. Kahn, who believes that wealth and technology will solve most problems, presents an assortment of visions of the future. Man’s impact on the fauna in the Pacific biota is presented in a scholarly account by lan McTaggart Cowan, the President of the Pacific Science Associa- tion. He examines technological development in relation to the disappearance of many species of fauna Vol. 91 and appeals to man’s imagination to save the ecosystems. J. D. Issacs and P. A. Larkin write about a much neglected subject, science policy. Both are aquatic biologists and both review the philosophy behind decision-making and policy formulation in the region. Issacs is unhappy with the present system because it represents the contributions of too few countries. Larkin feels that man’s technological development is progressing more rapidly than is socially desirable, and that society is incapable of developing a com- petent policy under such circumstances. The last essay in the book is by Thor Heyerdahl, a well known anthropologist and archaeologist, whose adventures in primitive crafts have lead him to alter many theories of early navigation. He writes about primitive navigation and simultaneously reveals an interesting and factual story about native cultures. I have refrained from making comments about Frank Fenner’s essay, “Options for Man’s Future: A Biologists View,” because it deserves special mention. Essentially, it is a capsule summary of the major issues surrounding man in the Pacific, and in fact, the world. But more than that, the essay is a well documented account, with illustrative graphs and charts, of man’s changing position on earth in relation to the environ- ment in which he lives. It is recommended that this essay be read before any of the others. This book has a conscience. It is an excellent addition to the continuing saga of world dynamics. PAUL A. GRAY 16 Cavehill Crescent, Scarborough, Ontario MIR 4P9 Erosion of Land in Northwestern Alberta — Report and recommendations By W.R. Trost (Chairman). 1976. Alberta Environment Conservation Authority, Edmonton. 73 pp. Free. The report on Erosion of Land in Northwestern Alberta by the Alberta Environment Authority was instigated as a result of the interest generated by the Peace River Regional Agricultural Service Board (1974), the Provincial Conference of Agricultural Service Boards (February 1975), and to the resultant public petition of 200 signatures. The area which the report encompasses 1s west and north from the city of Edmonton to the provincial boundaries with British Columbia and the Northwest Territories, or roughly an area of 100 000 square miles. In this region two areas were under study: the Peace River District and Lesser Slave Lake Watershed. It was felt that each had its inherent erosion control problems. Recommendations on erosion control were to be made for each respective district. The region had two major periods of settlement, the first of which was at the time of the completion of a railroad to the area in 1916. The second settlement was at the advent of the Second World War and the completion of the Alaskan Highway. The first exploiters of the land were agriculturalists, but the discovery of varied mineral resources led to the diversification of land utilization. It became apparent to the early settlers that an erosion problem existed. It was found that the causes of some of the problem were due to the adaptation of an inappropriate Great Plains farming technology to Northwestern Alberta. Government and public in- terest was stimulated to examine ways in which they could, along with the educational institutions and land managers, pool their collective resources to accom- 1977 plish (1) solutions to the problem of land reclamation due to past erosion difficulties, and (2) the prevention of future losses due to agricultural or other breakings. Alberta will have to live with past management decisions but this report is indicative of a new trend for making management decisions on new frontiers. The basic premise underlying the report is that every frontier is characteristic in nature of ‘geological, hydrological, and vegetational properties. Manage- ment decisions affecting the utilization of any one of these resources will have to be considered in the context of each individual property as it affects the total situation. A number of government and private institutions have jurisdiction over each property which affects the outcome of land use. Therefore it is imperative that land management decisions be inte- grated with each jurisdiction prior to the establish- ment of a land policy. This is to be done witha referral system situated in government to deal with the total problem through consultation with other departments and agencies affected. All relevant factors can thus be considered and integrated solutions can be achieved. Northwestern Alberta has 5 million acres presently being farmed. The potential exists to increase farm acreage by 10-15 million acres in addition to a potential of 50 million acres for forestry production. The area is, in effect, a new frontier. The government realizes the fragility of the region and in order to ensure its continued productivity and minimize erosion losses, there is an increased interest in developing policies for proper resource and land utilization. Although the subject matter of the work may not be NEW TITLES Zoology African ungulates. A comparative review of their ethology and behavioral ecology. 1977. By Walter Leuthold. Volume 8. Zoophysiology and Ecology. Springer-Verlag, New York. xiv + 308 pp., illus. $31.70. The Audubon Society book of wild birds. 1976. By Les Line and Franklin Russell. Abrams, New York. 292 pp., illus. $35. The biology of sticklebacks. 1976. By R.J. Wootton. Academic, New York. x + 385 pp., illus. $29.50. The book of turtles. 1977. By R. E. Nicholls. Running Press, Philadelphia. 152 pp., illus. Paper $4.95. Collected papers in avian paleontology honoring the 90th birthday of Alexander Wetmore. 1976. Edited by Storrs L. Olson. Smithsonian Contributions to Paleobiology Number 27. Smithsonian Institute Press, Washington. xxvi + 212 pp., illus. BOOK REVIEWS 433 of interest to all, the manner in which the government proceeded to do the report is indicative of a newera of decision-making. The report takes into consideration the needs of the land manager, the diversity of government jurisdictions, and the characteristics of the region under scrutiny. All facets have an input into the decision-making process. The Albertan Department of Transport’s referral system was recommended as a model for the Depart- ment of Agriculture and others in soil surface operations. However, it was stated in passing in the report. It would have been useful to have included a practical example of how the system works. The work is rather choppy at times but this is owing to the large number of briefs presented to the Authority. To try to represent all points of view in 70 pages 1s at best a monumental task. An important underlying principle of the report was that of public input. In this writer’s opinion the work was weighted in the realm of government and professional testimony. Perhaps the situation would have been better clarified had the report included a model of government-professional-public interaction in the decision-making process so that the reader could determine how a balance could be struck. Overall I believe the work will be of interest and of value to all persons involved in land management and planning. GERRY MADIGAN Woodland Resources, Macdonald College, Ste Anne de Bellevue, Quebec HOA 1CO Bears. Their biology and management. 1976. Edited by M. R. Pelton, J. W. Lentfer, and G. E. Folk. Papers froma conference, Binghampton, New York, June 1974 and a congress, Moscow, June 1974. IUCN Publications New Series Number 40. International Union for Conservation of Nature and Natural Resources, Morges, Switzerland. 468 pp., illus. Paper $12. Eagles of the world. 1977. By Leslie Brown. Universe, New York. 224 pp., illus. $12.50. +Handbuch der Voegel Mitteleuropas. Band 7. Charadrii- formes (2. Teil). 1977. Edited by U. N. G. von Blotzheim, K. M. Bauer, and E. Bezzel. Akademische, Wiesbaden. 893 pp., illus. +How reptiles and amphibians live. 1977. By A. C. Ech- ternacht. Volume 6 of How animals live. Edited by P. Hutchinson. Elsevier Phaidon, Oxford. (Canadian distri- 434 THE CANADIAN FIELD—NATURALIST butor Burns and MacEachern, Don Mills). 142 pp., illus. $12.50. An illustrated key to freshwater and soil amoebae with notes on cultivation and ecology. 1976. By F. C. Page. Scientific Publication Number 34. Freshwater Biological Association, Ambleside, England. 156 pp. Paper £2.50. Lambert’s birds of garden and woodland. 1977. By A. Mitchell. Paintings by T. Lambert. Scribner, New York. 128 pp. $12.50. Little mammals of the Pacific Northwest. 1977. By Ellen. Krilzman. Douglas, Douglas, David, and Charles, Van- couver. 128 pp., illus. Paper $5.95. Lost wild worlds: the story of extinct and vanishing wildlife of the eastern hemisphere. 1976. By R.M. McClung. Morrow, New York. 288 pp., illus. $8.95. Murex shells of the world. An illustrated guide-to the Muricidae. 1976. By G. E. Radwinand A. D’Attilio. Color photographs by D. K. Mulliner. Stanford University Press, Stanford. xii + 284 pp. + plates. $35. The polychaete worms. Definitions and keys to the orders, families, and genera. 1977. By K. Fauchald. Natural History Museum of Los Angeles County, Los Angeles. x + 188 pp., illus. Paper $7. {Proceedings of the 1975 predator symposium. !977. Edited by Robert L. Phillips and Charles Jonkel. Montana Forest and Conservation Experimental Station, University of Montana, Missoula. ix + 268 pp., illus. Paper $4. Reptile ecology. 1976. By Harold Heatwole. Australian Ecology Series. University of Queensland Press, New York. xvill + 178 pp., illus. Cloth $20; paper $9.95. *Rocky Mountain wildlife. Ecology, behavior, identification, distribution. 1976. By Don Blood, Tom W. Hall, and Susan I. Baumgarten. Hancock House, Saanichton, British Columbia. 132 pp., illus. }Running, walking and jumping. The science of locomotion. 1977. By Anne Innis Dagg. Wykeham, London. x + 143 pp., illus. Paper £3.25; cloth £5.75. Shells and shores of Texas. 1977. By Jean Andrews. Uni- versity of Texas Press, Austin. xx + 366 pp., illus. $19.95. *Snakes — a natural history. 1977. By Parker and Grandison. Cornell University Press, Ithaca. Cloth $8.95; paper $3.95. Theories of populations in biological communities. 1977. By F. B. Christiansen and T. M. Fenchel. Ecological Studies Volume 20. Springer-Verlag, New York. x + 144 pp., illus. $27.30. Botany Defense mechanisms of plants. 1977. By B. J. Deverall. Vol. 91 Monographs in Experimental Biology Number 19. Cam- bridge University Press, New York. viii + 110 pp., illus. $12.50. The flora of Manitoulin Island and the adjacent islands of Lake Huron, Georgian Bay and the North Channel. 1977. By J.K. Morton. University of Waterloo Biology Series Number 15, Waterloo. 62 pp. $4.50. Fossil algae. Recent results and developments. 1977. Edited by Erik Fluegel. Papers from a symposium, Erlanger, West Germany, October 1975. Springer-Verlag, New York. xiv + 376 pp., illus. $36.10. { Genera of the eastern plants. A guide to the genera of native and commonly introduced ferns and seed plants of eastern North America from the Atlantic to the Great Plains from Key West-Southern Texas into the Arctic. 1977. By Wade T. Batson. 3rd edition. Wiley, New York, Toronto. 203 pp., illus. Paper $5.50. Hortus third. A concise dictionary of plants cultivated in the United States and Canada. 1977. Initially compiled by L. H. and E. Z. Bailey. Revised and expanded by the staff of the Liberty Hyde Bailey Hortorium. MacMillan, New York. 1290 pp. $99.50. Northland wildflowers. A guide for the Minnesota region. 1977. By J.B. Moyle and E. W. Moyle. University of Minnesota Press, Minneapolis. x + 236 pp., illus. $12.95. A plant geography of Alberta. An interpretation based on the 1965 vegetation map. 1976. By Margaret E. E. North. Studies in Geography Monograph 2. University of Alberta Department of Geography, Edmonton. xiv + 148 pp. + map. Paper $5. Seaweeds. A color-coded, illustrated guide to common marine plants of the east coast of the United States. 1977. By C. J. Hillson. Pennsylvania State University Press, Univer- sity Park. x + 194 pp. Cloth $10; paper $6.95. Who named the daisy? Who named the rose? A roving dictionary of North American wild flowers. 1977. By Mary Durant. Drawings by Eleanor Cooney. Dodd, Mead, New York. x + 214 pp. $7.95. Wildflowers of Manning Park. 1976. By J. L. Underhill and C. C. Chuang. British Columbia Museum, and British Columbia Provincial Park, Victoria. 144 pp. Environment America’s great outdoors. The story of the eternal romance between man and nature. 1977. Edited by L. J. Bashline and D. Saults. Ferguson, Chicago. xvi + 368 pp., illus. + plates. $24.95. Conservation of resources. 1977. Papers from a symposium, Glasgow, April 1976. Special Publication Number 27. Chemical Society, London. x + 246 pp., illus. Paper $12. Ecology and the politics of scarcity. Prologue to a political 1977 theory of the steady state. 1977. By William Ophuls. Freeman, San Francisco. xiv + 304 pp. Cloth $12.95; paper $6.95. Ecology out of joint. New environments and why they happen. 1977. By L. J. and M. Milne. Scribner, New York. xi + 304 pp., illus. $8.95. Ecosystem research in South America. 1977. Edited by Paul Muller. Biogeographica Volume 8. Junk, The Hague. vi + 152 pp., illus. Dfl. 45. Evolution and ecology. Essays on social transformation. 1977. Edited by J. H. Steward, J. C. Steward, and R. F. Murphy. University of Illinois Press, Urbana. x + 406 pp. $12.95. Marine ecology. Selected readings. 1976. Edited by J. S. Cobb and M. M. Harlin. University Park Press, Baltimore. xii + 546 pp., illus. Paper $16.50. Muskeg and the northern environment in Canada. 1977. Edited by N. W. Radforth and C. O. Brawner. University of Toronto Press, Toronto. 399 pp., illus. $35. +A nature conservation review. The selection of biological sites of national importance to nature conservation in Britain. Volumes I and II. 1977. Edited by Derek Ratcliffe. Cambridge University Press, London, New York. xvi + 401 pp. + maps, vill + 319 pp. $69.50, $49.50. Outdoors Canada. A unique and practical guide to our wilderness and wildlife. 1977. Edited by Douglas R. Long. Canadian Automobile Association and Reader’s Digest, Montreal. 384 pp., illus. $21.77 (to members). Seagrass ecosystems. A scientific perspective. 1977. Edited by C. P. McRoy and C. Helferich. Papers from a workshop, Leiden, Netherlands, October 1973. Marine Science Volume 4. Dekker, New York. xi + 314 pp., illus. $29.75. The swamp. 1976. By Bill Thomas. Norton, New York. 223 pp., illus. $24.95. Underwater life: the world you never see. 1976. By P. Parks. Rand McNally, Chicago. 128 pp., illus. $9.95. Wet coastal ecosystems. 1977. Edited by V. J. Chapman. Ecosystems of the world. Volume 1. Elsevier, New York. xii + 428 pp., illus. $49.95. BOOK REVIEWS 435 Other +The aquatic explorers: a history of the Fisheries Research Board of Canada. 1977. By Kenneth Johnstone. University of Toronto Press, Toronto. xv + 342 pp., illus. $20. The collected papers of Charles Darwin. 1977. Edited by P.H. Barrett. University of Chicago Press, Chicago. 2 volumes. xvill + 278 pp., illus. and viii + 326 pp. $20 each. Field guide to snow crystals. 1977. By E. R. LaChapelle. Douglas, Douglas, David, and Charles, Vancouver. 104 pp., illus. Paper $4.95. Interpreting the earth. 1977. By Robert R. Compton. Harcourt Brace Javanovich, New York. xx + 554 pp.., illus. $14.95. Natural resources of British Columbia and the Yukon Territory. 1977. By Mary L. Barker. Douglas, Douglas, David, and Charles, Vancouver. 166 pp., illus. $13.95. Planet earth: encyclopedia of geology. 1977. Edited by A. Hallam. Plays (Canadian distributor Burns and Mac- Eachern, Toronto). 320 pp., illus. $24. +Point Pelee. Canada’s deep south. 1977. By Darryl] Stewart. Burns and MacEachern, Toronto. 112 pp., illus. Paper $5.95. +Science Council of Canada study on population, technology and resources. Perceptions 4, people and agricultural land. 1977. By Charles Beaubien and Ruth Tabacnik. Science Council of Canada, Ottawa. 137 pp., illus. Canada $4; elsewhere $4.80. Soils of the polar landscapes. 1977. By John C. F. Tedrow. Rutgers University Press, New Brunswick, New Jersey. xxvi + 638 pp., illus. $60. + Weather almanac. A reference guide to weather, climate and air quality in the United States and its key cities comprising statistics, principles and terminology. 1977. Edited by J. A. Ruffner and F. E. Blair. 2nd edition. Gales, Detroit. 728 pp. $25. tavailable for review *assigned for review Index to Volume 91 Compiled by R. Emerson Whiting Abies balsamea, 404, 414 spp., 369 Abraham, K.F., P. Mineau, and F. Cook. Unusual predators of Snow Goose eggs, 317 Abutilon theophrasti, 359 Acanthis flammea, 317 Acarospora fuscata, 307 Acer, 424 macrophyllum, 309 nigrum, 388 rubrum, 405, 414 Achillea millifolium, 368 Acris creptians blanchardi, 385 Actaea pachypoda, 378 Actitis macularia, 227 Adams, W. J., 294 Aeshna canadensis, 98 Age, juvenile, at emergence and Gestation period in Richardson’s ground squirrel, 410 Agropyron violaceum, 12, violaceum spp. andinum, 320 Agroeca ornata, 402 Alaska, The river otter (Lutra canadensis) on the north slope of the Brooks Range, 303 Alaskan Rosa acicularis, Germination requirements of, 58 Alaskan vascular flora, New and notable finds in the, 319 Alberta, First record of Anna’s Hummingbird for, 394 Alberta, First report of the tiger trout hybrid, Salmo trutta Linnaeus X Salvelinus fontinalis (Mitchill), in, 85. Alberta, Nesting biology of the Sora at Vermillion, 63 Alberta, northeastern, Northern range extension for the brassy minnow in, 402 Alberta, Range of the bushy-tailed wood rat (Neotoma cinerea) in, 323 Alberta, southwestern, Summer food habits of Golden Eagles in, 296 Alberta, southwestern, Summer habitat use by White-tailed Ptarmigan in, 367 Alces alces, 32, 94, 96 Alces alces, Population fluctuations of wapiti (Cervus elaphus) and moose, in Riding Mountain National Park, Manitoba, 1950-1976, 130 Alectoria vancouverensis, 308 Aleksiuk, M. Sources of mortality in concentrated garter snake populations, 70 Alex, J. F. (reviews by), 204, 205 Alisma plantago-aquatica, 64 Allan, J. H. First report of the tiger trout hybrid, Salmo trutta Linnaeus X Salvelinus fontinalis (Mitchill), in Alberta, 85 Allium cernuum, 389 Allolobophora chloratica, 395 Alnus, 365 crispa, 242 rubra, 308 rugosa, 404 Alopecurus alpinus, 14, 82 Alopex lagopus, 15 Amaurobius borealis, 402 Ambloplites rupretis, 425 Ambrosia psilostrachya, 359 Ambystoma texanum, 385 Ammannia coccinea, 384 Amphipoda: Gammaridae, The genus Crangonyx in the Central Connecticut River system, 256 Anaphalis margaritacea, 378 Anarhichas denticulatus, The wolffish, cf., new to the Amundsen Gulf area, Northwest Territories, and a probable prey of the ringed seal, 288 Anas acuta, 316 platyrhynchos, 32, 286 rubripes, | strepera, 248 Andropogon scoparius, 350 virginicus, 350 Androsace chamaejasme, 368 Anemone drummondi, 368 Animal Behavior Society, Northeastern regional meeting of the, 102 Anser caerulescens, 316 c. caerulescens, 317 Antennaria alpina var. media, 368 lanata, 368 spp., 367 monocephala ssp. monocephala, 320 spp. philonipha, 322 Anthus spinoletta, 317 Antilocarpa americana, 282 Aporrectodea trapezoides, 395 tuberculata, 395, 360 turgida, 395 Aquila chrysaetos, 296 Aralia hispida, 378 nudicaulis, 378, 414 Arcagrostis latifolia, 12 Arctophila fulva, 319 Ardea herodias, 88 Arenaria chamissonis, 320 interpres, 227 sajanensis, 368 Argus, G. W., review by, 107 Aristida basiramea, 350 dichotoma, 350 gracilis, 351 intermedia, 351 longispica, 350 necopina, 350 oligantha, 350 purpurascens, 350 Arnica alpina, 82 cordifolia, 368 Arthopyrenia halodytes, 307 Arthronia radiata, 308 1977 Arthrothelium cfr. ilicinum, 308 Aspicilia cinerea, 307 Asplenium platyneuron, 84, 265 trichomanes, 84, 265 viride, 267 Aster acuminatus, 378 azureus, 357 macrophyllus, 414 Astragalus alpinus, 82 occidentalis, 368 Ataenius strigatus, 196 Athyrium filix-femina, 265, 378, 405 Pycnocarpon, 265 thelypterioides, 265 Atlantic, north, The changing sea-bird populations of the, 102 Atrichum crispum, 405 Authors and references, Note to, 101 Avifauna of the West Foxe Islands, Northwest Territories, 1956-1976, Changes in the, 314 Avocet, American, 230 Axtell, H. H., P. Benham and J. E. Black. Spotted Red- shank sighted in southern Ontario, 90 Axyrus amaranthoides, 251 Aythya affinis, 248, 286 americana, 248 Bacidia friesiana, 308 Baird, D. J., 177 Banfield, A. W. F., reviews by, 200, 201, 203, 329 Barkhouse, P., 377 Barrett, M. W., 94 Bartramia longicauda, 227 Basiaeschna janata, 98 Bass, rock, 425 smallmouth, 425 Bear, 347 black, 96, 318 Bears, nuisance black, (Ursus americanus) in southeastern British Columbia, Movements of, 419 Beaver, 32, 365 Beckmannia syzigachne, 164 Bec-scie, common, | Behavior, Manipulative, by a red squirrel, 417 Benham, P., 90 Benton, F., I. M. Brodo, and D. H. S. Richardson, Lichens of the Bamfield Marine Station, Vancouver Island, British Columbia, 305 Berrill, M., 406 Berry, D.K. Northern range extension for the brassy minnow in northeastern Alberta, 402 Betula glandulosa, 246 Papyrifera, 413 sp., 277 Bidens connata, 388 Bider, J. R., 141 Biological flora of Canada, The — a new series, 269 Bird, D. M. and J. Wright. Apparent distraction display by a Barred Owl, 176 Bison bison, 418 Bison, Harassment of an elk calf by, 418 Black, J. E., 90 Blarina brevicauda, 43, 180 INDEX TO VOLUME 91 437 Boag, D. A. Summer food habits of Golden Eagles in southwestern Alberta, 296 Bobcat (Lynx rufus) specimens from southern Ontario, Two recent, 98 Boles, B. K. Predation by wolves on wolverines, 68 Bombycilla cedrorum, 191 garrula, 191 Bonasa umbellus, 297 Botrychium dissectum f. dissectum, 264 dissectum f. obliquum, 264 dissectum var. oneidense, 264 lanceolatum, 267 matricariaefolium, 264 multifidum, 264 simplex var. laxifolium, 264 simplex var. simplex, 264 simplex var. tenebrosum, 26 ternatum, 267 Y virginianum, 264 Bourget, A., | Boyd, H., review by, 105 Boyle, N. G. H., 322 Bradstreet, M.S. W., G. W. Page, and W.G. Johnston, Shorebirds at Long Point, Lake Erie, 1966-1971: seasonal occurrence, habitat preference, and variation in abundance, 225 Brant, Atlantic, 315 Branta bernicula hrota, 315 canadensis hutchinsti, 315 c. interior, 315 British Columbia, 189, 193 British Columbia, First record of Atka mackerel, Plewro- gammus monopterygius (Hexagrammidae), in, 175 British Columbia, Lichens of the Bamfield Marine Station, Vancouver Island, 305 British Columbia, Records of the boreal toad from the Yukon and northern, !85 British Columbia, southeastern, Movements of nuisance black bears (Ursus americanus) in, 419 British Columbia, The Cattle Egret in, 87 Britton, D. M., 262 Britton, D. M. The fern Woodsia obtusa (Spreng.) Torrey in Ontario, 84 Brodo, I. M., 305 Bromus inermis, 251 japonicus, 388 pumpellianus, 320 Browning, M. R. Geographic variation in Dunlins, Calidris alpina, of North America, 391 Brunton, D. F., 422 Bryoria trichodes, 308 Bubulcus ibis, 87 Bucephala albeola, 3 clangula, |, 286 islandica, | Buech, R.R., R.M. Timm, and K. Siderits. A second population of rock voles, Microtus chrotorrhinus, in Minnesota with comments on habitat, 413 Buellia grisovirens, 305 penichra, 308 punctata, 308 stellulata, 307 438 stillingiana, 308 Bufflehead, 3 Bufo americanus, 322 boreas boreas, 185 Bunting, Snow, 317 Buteo albonotatus, 310 Butler, R. W., 189 Butomus umbellatus, 388 By-laws, The Ottawa Field-Naturalists’ Club, 114 By-laws of the Ottawa Field-Naturalists’ Club, Notice of change to the, 325 Cactus, Prairie fires and pronghorn use of, 282 Calamagrostis canadensis, 72 spp., 96 Calcarius lapponicus, 191, 317 ornatus, 191 Calcium subquercinum, 308 Calidris alpina, 227, 391 a. arcticola, 391 a. hudsonia, 391 a. pacifica, 391 a. sakhalina, 391 bairdii, 227 canutus, 230 fuscicollis, 230 maritima, 230, 316 mauri, 230 melanotos, 227 minutilla, 227 pusilla, 227, 316 Calidris alpina, Geographic variation in Dunlins of North America, 391 Callioplus euoplus, 402 Caloplaca cerina var. chlorina, 307 ferruginea, 309 marina, 307 scopularis, 307 sp., 307 thallincola, 305 Caltha palustris, 82 Calypte anna, 394 Camassia scilloides, 384 Campbell, C. A. and A. A. Reznicek. New vascular plant records on Pelee and East Sister Islands, Essex County, Ontario, 384 Campbell, C. A. and D. M. Britton. Pteridophytes of the Regional Municipality of Waterloo, 262 Campbell, C. A., review by, 427 Campbell, R. W. Use of man-made structures as nest sites by Pigeon Guillemots, 193 Campbell, R. W. and W.C. Weber. The Cattle Egret in British Columbia, 87 Camptosorus rhizophyllus, 265 Canada, A new series on the biological flora of, in The Canadian Field-Naturlaist, 101 Canada, Distribution and abundance of The Prairie Rattlesnake, Crotalus viridis viridis, in, 122 Canada, Distribution of the auricled twayblade orchid (Listera auriculata) in, and description of new stations in southern Ontario, 403 Canada, Genetic variants in, of the rainbow trout, Sa/mo gairdneri, called golden trout and palomino trout, 93 THE CANADIAN FIELD-NATURALIST Vol. 91 Canada, Paspalum ciliatifolium, a grass new to, from southwestern Ontario, 422 Canada, Pilot study for a Biological Survey of The Insects of, 199 Canada, The biological flora of — a new series, 269 Canadian Field-Naturalist, A new series on the biological flora of Canada in The, 101 Canard kakawi, | noir, | Canis lupus, 15, 68, 77, 96, 323 I. arctos, 77 “Cannibalism” and “scavenging” in ecological literature, Usage of the terms, 416 Capella gallinago, 227 Cardamine bellidifolia, 82 Carduelis tristis, Increase in overwintering by the American Goldfinch, in Ontario, 165 Carex atheroides, 64, 372 crinita, 405 davisii, 384 divulsa subsp. leersii, 384 gracillima, 405 intumescens, 405 misandra, 82 nardina, 14 peckii, 320 rugosperma, 351 sartwellii, 388 spp., 14, 63, 72, 96, 251, 367 stans, 14, 82, 321 Caribou, 17, 81, 96, 317 Peary, 78 Caribou, mountain, Summer use of a highway crossing by, 312) Carpenter, J. W. and C. G. Paterson. Variation in selected meristic series in the golden shiner, Notemigonus crysoleucas (Mitchill), in the New Brunswick — Nova Scotia border region, 74 Carya ovalis, 388 ovata, 85 Cassiope tetragona, 82, 367 Castilleja occidentalis, 368 Castor canadensis, 32, 365 Catling, P. M., 403 Catling, P. M., A. A. Reznicek, and J. L. Riley. Some new and interesting grass records from southern Ontario, 350 Catling, P. M. and W. Freedman. Melanistic Butler’s garter snakes (Thamnophis butleri) at Amherstburg, Ontario, 397 Catoptrophorus semipalmatus, 230 Cattle, 32, 63, 87 Caves, Terrestrial Oligochaeta of some New Brunswick, with remarks on their ecology, 360 Celtis tenuifolia, 384 var. soperi, 386 Cepphus columba, 193 grylle, 3 g. ultimus, 317 Cerastium arcticus, 82 Cervus canadensis, 95 elaphus, 418 1977 Cervus elaphus, Population fluctuations of wapiti, and moose (Alces alces) in Riding Mountain National Park, 1950-1976, 130 Cetrelia cetrarioides, 308 Chaenorrhinum minus, 356 Chaenotheca brunneola, 308 Chaerophyllum procumbens, 384 var. procumbens, 389 var. shortii, 389 Chalmers, G. A. and N. W. Barrett. paralysis in a moose calf, 94 Chapdelaine, G. Le Grand Cormoran (Phalacrocorax carbo) en hiver, le long des cétes de la Péninsule de Gaspésie, Québec, 184 Charadrius melodus, 227 semipalmatus, 91, 227, 316 vociferus, 227 Chenopodium album, 359 foggii, 384 Chevendra, C. R., reviews by, 208, 208, 335, 336, 336 Chicken, 32, 87 Chipmunk, eastern, 180 least, 41 Chrysanthemum bipinnatum ssp. bipinnatum, 322 Chrysemys picta, 47 p. bellii, 48 p. dorsalis, 55 Chrysemys picta marginata (Agassiz), Morphological par- ameters and spring activities in a central Ontario population of midland painted turtle, 47 Chrysosplenium tetrandum, 82 Cirsium arvense, 251 Cladonia chlorophaea, 308 macilenta subsp. theiophila, 308 mitis, 246 scabriuscula, 308 subsquamosa, 308 transcendens, 308 Clangula hyemalis, 1, 316 Clethrionomys gapperi, 41, 173, 178 rutilis, 96, 239, 417 Clethrionomys rutilus, Incidence of the dark color phase in tundra and taiga populations of northern red-backed voles, 173 Climacium dendroides, 405 Clintonia borealis, 178, 378, 414 Cochlearia officinalis, 82 Cody, W.J., reviews by, 206, 206, 207, 207, 334, 334 Colaptes auratus, 192, 297 Colloquium, Third annual Ontario ecological, 102 Colombie-Brittanique, Le ver québécois (Lumbricus festi- vus) envahit la, 395 Colonial Waterbird Group first annual meeting, 199 Color-banded semipalmated and Least Sandpipers, Request for information, 198 Colpodium Vahlianum, 14, 319 Coluber constrictor foxi, 385 Commelina communis, 388 Comments and policy, Editor’s, 221 Concerns, Environmental, 101 Connecticut River system, central, The genus Crangonyx (Amphipoda: Gammaridae) in the, 256 Trauma-induced INDEX TO VOLUME 91 439 Conobea multifida, 384 Cooch, F. G. Changes in the avifauna of the West Foxe Islands, Northwest Territories, 1956-1976, 314 Cook anes Cook, F. R. Records of the boreal toad from the Yukon and northern British Columbia, 185 Coptis trifolia, 378 Cordulia shurtleffi, 97 Cormoran, Le Grand, (Phalacrocorax carbo) en hiver, le long des cétes de la Péninsule de Gaspésie, Québec, 184 Cormorant, Great, 3 Cornus canadensis, 178, 378, 414 racemosa, 357 rugosa, 414 Corvus brachyrhynchos, 65, 70, 248 corax, 32 Corydalis flavula, 384 Corylus cornuta, 414 Cottus cognatus, Occurrence of the slimy sculpin in the Missouri drainage system, 415 Cougar in Manitoba, Status and habits of the, 28 Coutts, R. A., 322 Cowbird, Brown-headed, 346 Craig, R. E.and E. J. Crossman. Genetic variants in Canada of the rainbow trout, Salmo gairdneri, called golden trout and palomino trout, 93 Crangonyx (Amphipoda: Gammaridae) in the central Con- necticut River system, The genus, 256 Crangonyx pseudogracilis, 256 richmondensis, 256 r. laurentianus, 256 r. richmondensis, 256 Crepis nana, 320 Crins, W. J., P. D. Pratt, and D. F. Brunton. Paspalum ciliatifolium, a grass new to Canada from southwestern Ontario, 422 Crocethia alba, 227 Croskery, P., reviews by, 106, 207, 327, 328 Crossman, E. J., 93 Crotalus viridis viridis, Distribution and abundance of the Prairie Rattlesnake in Canada, 122 Crow, 65, 70, 248 Crypsis schoenoides, 358 Cryptogramma stelleri, 267 Cycloloma atripicifolium, 388 Cydonia oblonga, 384 Cymatogaster aggregata, 88 Cyperus erythrorhizos, 388 odoratus, 358 var. squarrosus, 389 spp. 389 Cypripedium acaule, 378 Cystopteris bulbifera, 265 fragilis, 84 fragilis var. mackayi, 265 protusa, 266 Dagg, A. I., reviews by, 106, 203, 3 Danthonia spicata, 351 Deer, Formosan, 32 mule, 28, 297 white-tailed, 28 Delphinium bicolor, 368 w ws) ies) Ww es) Ww — ao ~~ ~~ BN i) Ne) 440 THE CANADIAN FIELD-NATURALIST Vol. 91 Dendragapus obscurus, 297 X triploidea, 266 Dendrobaena octaedra, 395 X uliginosa, 266 Dendrodrilus rubidus, 360, 395 Duck, 291 Dendroica coronata, 317 Black, | Dennstaedtia punctilobula, 265 Dunlin, 227 Densmore, R. and J. C. Zasada. Germination requirements Dunlins, Calidris alpina, of North America, Geographic of Alaskan Rosa acicularis, 58 variation in, 391 Deschampsia brevifolia, 14 Dupontia fisheri, 319 cespitosa var. parviflora, 350 : Dyer, M., review by, 428 Design for a small mammal live trap, An improved, 399 Eagles, Golden, in southwestern Alberta, Summer food Desmognathus fuscus, 299 habits of, 296 Didymops transversa, 98 Eclipta prostrata, 384 Diervilla lonicera, 414 Ecological literature, Usage of the terms “cannibalism” Dimerella lutea, 308 and “scavenging” in, 416 Diplachne acuminata, 353 Editorial policy, 117 Diplotaxis muralis, 388 Editor's comments and policy, 221 tenuifolia, 388 Editor’s report, 197 Dog, 32, 69 Egret, Cattle, in British Columbia, the, 87 Douglasia gormanii, 320 Eider, Common, | Douglass, K.S., 72 Northern, 314 Douglass, R. J. and A. E. L. McNaughton. A recent record Eider, commun, | of the meadow jumping mouse, Zapus hudsonius, in Eisenia rosea, 360, 395 the Northwest Territories, 96 : Ejiseniella tetraedra, 396 Douglass, R. J.and K. S. Douglass. Micro-habitat selection Eleocharis sp., 65 of chestnut-cheeked voles (Microtus xanthognathus), Elk, 95 m2. american, 28 Douglass, R. J. Population dynamics, home ranges, and Elk calf, Harrassment of, by bison, 418 habitat associations of the yellow-cheeked vole, Empetrum nigrum, 246 Microtus xanthognathus, in the Northwest Territories, | Empidonax sp., 191 237 Environmental concerns, 101 Dowichers, 91, 225 Epilobium alpinum, 368 Long-billed, 226 angallidifolium, 320 Short-billed, 226 angustifolium, 378 Draba alpina, 82 davuricum var. arcticum, 322 fladnizensis, 321 _ latifolium, 14, 368 spp., 14 Epitheca cynosura, 97 Dragonflies, Swarming of, noted at Drag Lake, Ontario, 97 princeps, 97 Dryas, 321 spinigera, 97 hookeriana, 367 Equisetum arvense, 263 integrifolia, 8, 82 arvense var. boreale, 263 Dryopteris assimilis, 266 fluviatile, 263 X boottii, 266 fluviatile X arvense, 267 X burgessii, 266 hyemale var. affine, 263 campyloptera, 266 hyemale var. affine X E. variegatum, 263 clintoniana, 266 laevigatum, 263 clintoniana X cristata, 266 laevigatum X E. variegatum, 263 clintoniana X intermedia, 266 limosum, 263 clintoniana X marginalis, 266 X litorale, 267 cristata, 266 X nelsonii, 263 cristata X intermedia, 266 palustre, 267. cristata X marginalis, 266 pratense, 263, 405 cristata x spinulosa, 266 — prealtum, 263 X dowellii, 266 scirpoides, 264, 368 goldiana, 266 spp., 72 intermedia, 266 sylvaticum, 264 intermedia X spinulosa, 266 X trachydon, 263 marginalis, 266 variegatum, 264 marginalis X spinulosa, 266 Eragrostis spectabilis, 350 phegopteris, 378 Eremophila alpestris, 191 X pittsfordensis, 266 Erethizon dorsatum, 33, 363 X slossonae, 266 Erigeron aureus, 368 spinulosa, 266, 378 grandiflorus, 368 spinulosa var. americana, 266 humulis, 368 1977 Erignathus barbatus, 181 Eriophorum angustifolium, 319 Scheuchzeri, 82 spp., 14 triste, 82, 321 Ermine, 96, 180, 239 Erskine, D., review by, 111 Essex County, Ontario, New vascular plant records on Pelee and East Sister Islands, 384 Euonymus atropurpureus, 388 Eupatorium altissimum, 384 Euphorbia corollata, 357 cyparissius, 388 dentata, 388 obtusata, 384 Euphrasia disjuncta, 321 Eurycea bislineata, 299 Eutamias minimus, 41, 180 Eutrema Edwardsii, 82 Fagus, 424 grandifolia, 176 Fahselt, D. Growth rates in vegetative and reproductive tissue of Parmelia cumberlandia after relocation to a new environment, 134 Falco columbarius, 190 spp. richardsonii, 192 Felis c. cougar, 38 c. hippolestes, 33 concolor missoulensis, 28 Fern Woodsia obtusa (Spreng.) Torrey in Ontario, The, 84 Ferron, J. and J. Prescott. Gestation, litter size, and number of litters of the red squirrel (Tamiasciurus hudsonicus) in Quebec, 83 Festuca baffinensis, 14 brachyphylla, 14 Fieldfare in Ontario, 91 Fillmore, E.R. and R.D. Titman. Chipping Sparrow hanged, 69 Fimbristylis autumnalis, 355 Fires, Prairie, and pronghorn use of cactus, 282 Fisher, 32 Flicker, Common, 192 Flora, Alaskan vascular, New and notable finds in the, 319 Flora, biological, of Canada, A new series on the, in The Canadian Field- Naturalist, 101 Flora of Canada, The biological — a new series, 269 Flycatcher, 191 Foottit, R.G. and R.W. Butler. Predation on nesting Glaucous-winged Gulls by river otter, 189 Fox, 69 arctic, 15 Fragaria virginiana, 368 Fraxinus americana, 85 quadrangulata, 389 Freedman, W., 397 Freitag, R., 401 Frog, 291 Blanchard’s cricket, 385 chorus, 20 cricket, 102 wood, 20, 322 Frog, mink, in the James Bay region of Quebec, Range extensions of the water shrew and, 322 INDEX TO VOLUME 91 441 Gadwall, 250 Galium triflorum, 378 Gall, J. M., 424 Gammaridae (Amphipoda), The genus Crangonyx in the central Connecticut River system, 256 Garrot, de Barrow, | commun, | Gaspésie, Québec, Le Grand Cormoran (Phalacrocorax carbo) en hiver, le long des cOtes de la Péninsule de, 184 Gaultheria hispidula, 378 procumbens, 378 Gavia arctica, 315 immer, 315 stellata, 315 Gentiana glauca, 368 Germination requirements of Alaskan Rosa acicularis, 58 Gestation period and juvenile age at emergence in Richard- son’s ground squirrel, 410 Geum Rossii, 82 Ginns, J., 194 Glaucomys volans, 424 Glyceria grandis, 64 striata, 405 Gnaphalium uliginosum, 388 Godfrey, Earl, Thank you, 198 Godin, J.-G. J. A Great Blue Heron preying on shiner perch in deep water, 88 Godwit, Hudsonian, 230 Goldeneye, 286 Barrow’s, | Common, | Goldfinch, American, Carduelis tristis, Increase in over- wintering by, in Ontario, 165 Gomphus spicatus, 98 Goose, Canada, 315 Lesser Snow, 316, 317 Goose, Snow, eggs, Unusual predators of, 317 Gorham, S. W. review by, 103 Goshawk, 188 Grackle, 65 Common, 346 Grackles, Common, Predatory behavior by, 187 Grand River system, Occurrence of the green sunfish (Leopmis cyanellus) in the, 424 Graphis elegans, 308 Grass new to Canada from southwestern Ontario, Paspalum ciliatifolium, a, 422 Grass records from southern Ontario, Some new and interesting, 350 Gray, P. A., review by, 431 Grebe, Horned, breeding habitat in Saskatchewan park- lands, 372 Gregory, P. T. Life history observations of three species of snakes in Manitoba, 19 Groundhog, 273 Grouse, 58 Growth rates in vegetative and reproductive tissue of Parmelia cumberlandia after relocation to a new environment, 134 Guillemot, Black, 3, 317 Guillemots, Pigeon, Use of man-made structures as nest 442 sites by, 193 Gull, Glaucous, 316 Great Black-backed, 314 Herring, 316 Kumlien’s, 316 Gull, Little, (Larus minutus) in Arctic North America, the, 294 Gulls, Glaucous-winged, Predation on nesting, by river otter, 189 Gulls, Herring, nesting on Brothers Island, Lake Ontario, in 1973, Reproductive success of, 148 Gulls, Wanted: data on the seasonal distribution of North American, 325 Gulo gulo, 68 Gymnocarpium dryopteris, 265 Gyrinophilus porphyriticus, 299 Habitat, Horned Grebe breeding, in Saskatchewan park- lands, 372 Habitat use by White-tailed Ptarmigan in southwestern Alberta, Summer, 367 Haemogamasus alaskensis, 179 ambulans, 179 Halliday, G. New and notable finds in the Alaskan vascular flora, 319 Hare, arctic (Lepus arcticus monstrabilis), on Axel Heiberg Island, Northwest Territories, Morphology, repro- duction, diet, and behavior of the, 8 Hare, Snowshoe, 32, 58 varying, 96 Harms, V. L., review by, 109 Hawk, Red-tailed, 188 Hawk, Zone-tailed, in Nova Scotia, A, 310 Heaney, L. R., 177 Hedwigia sp., 246 Hedysarum sulphurescens, 368 Heleochloa schoenoides, 358 Herbison, B., 419 Herbs, forest, and dwarf shrubs of the New Brunswick — Nova Scotia border region, The structure and rate of growth of the rhizomes of some, 377 Heron, Great Blue, preying on shiner perch in deep water, A, 88 Herpetology information needed, 102 Herzog, P.W. Summer habitat use by Ptarmigan in southwestern Alberta, 367 Heteranthera dubia, 388 Heterodon nasicus nasicus, 19 Hierochloe alpina, 14 Highway crossing by mountain caribou, Summer use of a, 312 Hines, J. E. Nesting and brood ecology of Lesser Scaup at Waterhen Marsh, Saskatchewan, 248 Hirundo rustica, 183 Hog, 32 Homo sapiens, 416 Hoplopsyllus glacialis glacialis, 14 Hordeum jubatum, 388 Horse, 32, 87 Humans, 416, 419 Hummingbird, Anna’s, First record of, for Alberta, 394 Humphreys, G. B., 424 Hussell, D. J. T. and M. J. Porter. Fieldfare in Ontario, 91 Hybognathus hankinsoni, 402 White-tailed THE CANADIAN FIELD-NATURALIST Vol. 91 Hydrophyllum appendiculatum, 389 virginianum, 388 Hylocichla guttata, 191 ustulata, 191 Hypericum ellipticum, 405 Hypogymnia enteromorpha, 308 Icmadophila ericetorum, 308 Illex verticillata, 405 IlIman, W. I., 194 Information, herpetology, needed, 102 Insects of Canada, Pilot Study for a Biological Survey of the, 199 International Shorebird Surveys, Request for participants, 197 Ipomea purpurea, 388 Tris pseudacorus, 388 Trodroprocne bicolor, 191 Ixodes angustus, 180 Jaeger, Parasitic, 316 Johnson, D. R. and M. C. Todd. Summer use of a highway crossing by mountain caribou, 312 Johnson, S.R. and W. J. Adams. The Little Gull(Larus minutus) in Arctic North America, 294 Johnston, W. G., 225 Junco, Dark-eyed, 191 Junco hyemalis, 191 Juncus gerardii, 357 sp., 65 Juniperus virginiana, 357 Killdeer, 227 Kingbird, 314 Eastern, 317 Kinglet, Ruby-crowned, 191 Knapton, R. W., 393 Knapton, R.W. Breeding status of Says Phoebe in Manitoba, 183 Knot, Red, 230 Kott, E. and G. B. Humphreys. Occurrence of the green sunfish (Lepomus cyanellus) in the Grand River system, 424 Krause, D. W. and B. G. Naylor. Range of the bushy-tailed wood rat (Neotoma cinerea) in Alberta, 323 La Roi, G. H. The biological flora of Canada — a new series, 269 Laelaps kochi, 179 Lagopus leucurus, 367 sp., 68 Lake Erie, Age and fecundity of the tadpole madtom, Noturus gyrinus, on Long Point, 292 Lake Erie, 1966-1971, Shorebirds at Long Point: seasonal occurrence, habitat preference, and variation in abundance, 225 Lake Ontario, Reproductive success of Herring Gulls nesting on Brothers Island, in 1973, 148 Lamium amplexicaule, 388 purpureum, 388 Larix laricina, 96 Lark, Horned, 191 Larus argentatus, 148 a. smithsonianus, 316 glaucescens, 189 glaucoides kumlieni, 316 marinus, 316 LOTT Larus minutus, The Little Gull in Arctic North America, 294 Lecanactis megaspora, 308 Lecania nylanderiana, 308 sp., 308 Lecanora campestris, 307 cfr. expallens, 308 fugiens, 307 muralis, 308 sp., 307 Ledum groenlandicum, 246 Lemming, bog, 41 northern bog, 239 southern bog, 180 Lepomus cyanellus, Occurrence of the green sunfish in the Grand River system, 424 Lepomis gibbosus, 425 macrochirus, 425 Lepraria candelaris, 308 incana, 308 Lepthyphantes zebra, 402 Leptochloa fasciscularis, 350 var. acuminata, 353 Leptogium palmatum, 308 Leptoloma cognatum, 350 Lepus americanus, 32, 96, 297 Lepus arcticus monstrabilis, on Axel Heiberg Island, Northwest Territories, Morphology, reproduction, diet, and behavior of the arctic hare, 8 Leucorrhinia frigida, 97 hudsonica, 97 Libellula julia, 97 quadrimaculata, 97 Lichens of the Bamfield Marine Station, Vancouver Island, British Columbia, 305 Ligusticum mutellinoides, 321 Limnodromus griseus, 226 scolopaceus, 226 sp., 91 spp., 225 Limosa haemastica, 230 Linaria dalmatica, 388 Lindernia dubia, 388 Lindsay, D. R. Weedy species at terminal grain elevators, Thunder Bay, Ontario, 158 Linnaea borealis, 246, 378, 414 Linum striatum, 355 Listera auriculata, Distribution of the auricled twayblade orchid in Canada and description of new stations in southern Ontario, 403 Listera convallarioides, 404 cordata, 404 spp., 403 Lobaria oregana, 309 pulmonaria, 308 spp., 307 Lobelia cardinalis, 388 Lobipes lobatus, 231 Long Point, Lake Erie, 1966-1971, Shorebirds at: seasonal occurrence, habitat preference, and variation in abundance, 225 Longspur, Chestnut-collared, 191 Lapland, 191, 317 INDEX TO VOLUME 91 443 Loon, Arctic, 315 Common, 315 Red-throated, 315 Lotus corniculatus, 388 Loucks, R. H. (review by), 209 Lowther, J. K. Nesting biology of the Sora at Vermillion, Alberta, 63 Ludwigia palustris, 388 Lumbricus festivus, Le ver québécois envahit la Columbie- Britannique, 395 Lumbricus rubellus, 395 terrestris, 360, 396 Lunaria annua, 388 Lutra canadensis, 189 Lutra canadensis, The river otter on the north slope of the Brooks Range, Alaska, 303 Luzula arctica, 14 confusa, 82 nivalis, 14, 82 Lycopodium annotinum, 263, 378 clavatum, 263, 378 complanatum var. flabelliforme, 263, 378 flabelliforme, 378 inundatum var. inundatum, 263 lucidulum, 263, 378 obscurum, 263, 378 obscurum var. dendroideum, 263 tristachyum, 267, 378 Lycopus uniflorus, 386 virginicus, 389 var. virginicus, 384 Lynx rufus gigas, 99 r. rugus, 99 r. superiorensis, 99 Lynx rufus, Two recent bobcat specimens from Southern Ontario, 98 Lysurus gardneri, an uncommon stinkhorn observed in eastern Ontario, 194 Lythrum alatum, 388 MacInnis, A., 310 Mackerel, Atka, Pleurogrammus monopterygius, (Hexa- grammidae), First record of, in British Columbia, 175 Macromia illinoiensis, 97 Madigan, G., reviews by, 335, 432 Madison, D. M. Movements and habitat use among inter- acting Peromyscus leucopus as revealed by radio- telemetry, 273 Madtom, tadpole, Noturus gyrinus, on Long Point, Lake Erie, Age and fecundity of the, 292 Magoun, A.J. and P. Valkenburg. The river otter (Lutra canadensis) on the north slope of the Brooks Range, Alaska, 303 Mahan, B.R. Harassment of an elk calf by bison, 418 Mahon, R. Age and fecundity of the tadpole madtom, Noturus gyrinus, on Long Point, Lake Erie, 292 Maianthemum canadense, 178, 378, 405 Mallard, 32, 286 Mammal, small, live trap, An improved design for a, 399 Mammal, small, populations, Changes in, after clearcutting of northern Ontario black spruce forest, 41 Man, 32, 70 Manitoba, Breeding status of Say’s Phoebe in, 183 Manitoba, Life history observations of three species of 444 snakes in, 19 Manitoba, Locations of winter dens utilized by striped skunks in Delta Marsh, 289 Manitoba, Population fluctuations of wapiti (Cervus ela- phus) and moose (Alces alces) in Riding Mountain National Park, 1950-1976, 130 Manitoba, Status and habits of the cougar in, 28 Marmota monax, 273 Martell, A. M. Incidence of the dark color phase in tundra and taiga populations of northern red-backed voles, Clethrionomys rutilus, 173 Martell, A. M. and A. Radvanyi. Changes in small mammal populations after clearcutting of northern Ontario black spruce forest, 41 Marten, 69, 96, 188 Martes americana, 69, 96 pennanti, 32 Matteuccia pensylvanica, 265 McAllister, D. E., reviews by, 330, 331 McAlpine, D.F. and J. W. Reynolds. Terrestrial Oli- gochaeta of some New Brunswick caves with remarks on their ecology, 360 McKeating, G.B. (ed.), Nature and urban man, 208 McLaren, I. A. and A. MacInnis. A Zone-tailed Hawk in Nova Scotia, 310 McNaughton, A. E. L., 96 MeNicholl, M. K. Usage of the terms “cannibalism” and “scavenging” in ecological literature, 416 McTaggart, S., 190 Meadowlark, Western, 191 Medeola virginiana, 378 Medicago falcata, 284 Meeting of the Animal Behavior Society, Northeastern regional, 102 Melandrium apetalum, 82 Melanerpes carolinus, 385 Melanitta perspicillata, 3 Melospiza lincolInii, 191 melodia, 191 Memogana, J., 181 Menegazzia terebrata, 308 Mephitis mephitis, 33, 71, 248, 289 Merganser, Common, | Red-breasted, 3 Mergus merganser, | serrator, 3 Merlins, Prey utilized by urban, 190 Micarea melaena, 308 Michener, G. Gestation period and juvenile age at emergence in Richardson’s ground squirrel, 410 Micropalama himantopus, 230 Micropterus dolomieui, 425 Mirosorex hoyi, 43, 96, 393 Microtus chrotorrhinus, 41 oeconomus, 417 pennsylvanicus, 41, 96, 178, 239, 274 sp., 70 xanthognathus, 96 Microtus chrotorrhinus, A second population of Rock Voles in Minnesota with comments on habitat, 413 Microtus chrotorrhinus, Natural history of rock voles in Minnesota, 177 Microtus xanthognathus, Population dynamics, home THE CANADIAN FIELD-NATURALIST Vol. 91 ranges,and habitat associations of the yellow-cheeked vole, in the Northwest Territories, 237 Middleton, A. L. A. Increase in overwintering by American Goldfinch, Carduelis tristis, in Ontario, 165 Middleton, A.L. A. Predatory behavior by Common Grackles, 187 Miller, F. R. and Russell, R. H. Unreliability of strip aerial surveys for estimating numbers of wolves on western Queen Elizabeth Islands, Northwest Territories, 77 Mineau, P., 317 Mink, 69, 96 Minnesota, A second population of rock voles, Microtus chrotorrhinus, in, with comments on habitat, 413 Minnesota, Natural History of rock voles (Microtus chrotor- rhinus) in, 177 Minnow, brassy, in northeastern Alberta, Northern range extension for the, 402 Minuartia biflora, 320 Missouri drainage system, Occurrence of the slimy sculpin, Cottus cognatus, in the, 415 Mitchella repens, 378 Monodon monoceros, Narwhals observed near King Chris- tian Island, Northwest Territories, 309 Monodon monoceros, The occurrence of a narwhal in Prince Albert Sound, western Victoria Island, Northwest Territories, 299 Monotropa uniflora, 378 Moose, 32, 96 Moose (Alces alces), Population fluctuations of wapiti (Cervus elaphus) and, in Riding Mountain National Park, Manitoba, 1950-1976, 130 Moose calf, Trauma-induced paralysis in a, 94 Mortality in concentrated garter snake populations, Sources of, 70 Morton, J. K., review by, 429 Morus rubra, 389 Mosquito, 317 Mouse, 70, 273, 401 deer, 41, 178, 363 meadow jumping, 239 Mouse, meadow jumping, Zapus hudsonius, in the North- west Territories, A recent record of the, 96 Muhlenbergia asperifolia, 350 schreberi, 388 uniflora, 355 Munroe, E., reviews by, 105, 326 Murphy, D., review by, 110 Murre, Thick-billed, 316 Muskox, 17, 78, 81 Muskrat, 96 Mustela erminea, 96, 180, 239 frenata, 297 vison, 69, 96, 180 Mutch, G. R. P. Locations of winter dens utilized by striped skunks in Delta Marsh, Manitoba, 289 Myosotis alpestris, 320, 368 macrosperma, 384 verna, 386 Myriophyllum spicatum, 388 Myth of the non-consumptive user, The, 343 Nagorsen, D. and R.L. Peterson. Two recent bobcat (Lynx rufus) specimens from southern Ontario, 98 Narwhal (Monodon monoceros) in Prince Albert Sound, 1977 western Victoria Island, Northwest Territories, The occurrence of a, 299 Narwhals (Monodon monoceros) observed near King Christian Island, Northwest Territories, 309 Naylor, B. G., 323 Nealis, V., J. Ginns and W.I. Illman. Lysurus gardneri, an uncommon stinkhorn observed in eastern Ontario, 194 Neotoma cinerea, 297 c. cinerea, 323 c. drummondii, 323 Neotoma cinerea, Range of the bushy-tailed wood rat in Alberta, 323 Neotrombicula harperi, 180 microti, 179, 413 Nephroma laevigatum, 308 Nero, R. W. and R. E. Wrigley. Status and habits of the cougar in Manitoba, 28 Nesting biology of the Sora at Vermillion, Alberta, 63 New Brunswick caves, Terrestrial Oligochaeta of some, with remarks on their ecology, 360 New Brunswick — Nova Scotia border region, The structure and rate of growth of the rhizomes of some forest herbs and dwarf shrubs of the, 377 New Brunswick — Nova Scotia border region, Variation in selected meristic series in the golden shiner, Note- migonus crysoleucas (Mitchill), in the, 74 North America, Arctic, The Little Gull (Larus minutus) in, 294 North America, Geographic variation in Dunlins, Calidris alpina, of, 391 North American Gulls, Wanted: data on the seasonal dis- tribution of, 325 Northwest Territories, A recent record of the meadow jumping mouse, Zapus hudsonius, in the, 96 Northwest Territories, Changes in the avifauna of the West Foxe Islands, 1956-1976, 314 Northwest Territories, Morphology, reproduction, diet, and behavior of the arctic hare (Lepus arcticus monstra- bilis) on Axel Heiberg Island, 8 Northwest Territories, Narwhals (Monodon monoceros) observed near King Christian Island, 309 Northwest Territories, Population dynamics, home ranges, and habitat associations of the yellow-cheeked vole, Microtus xanthognathus, in the, 237 Northwest Territories, The flowering phenology of common vascular plants at Bailey Point, Melville Island, 81 Northwest Territories, The occurrence of a narwhal (Mono- don monoceros) in Prince Albert Sound, western Victoria Island, 299 Northwest Territories, The wolffish, cf. Anarhichas denti- culatus, new to the Amundsen Gulf area, and a probable prey of the ringed seal, 288 Northwest Territories, Unreliability of strip aerial surveys for estimating numbers of wolves on western Queen Elizabeth Islands, 77 Notemigonus crysoleucas (Mitchill), Variation in selected meristic series in the golden shiner in the New Brunswick — Nova Scotia border region, 74 Noturus gyrinus, Age and fecundity of the tadpole madtom on Long Point, Lake Erie, 292 Nova Scotia, A Zone-tailed Hawk in, 310 Nova Scotia - New Brunswick border region, The structure INDEX TO VOLUME 91 445 and rate of growth of the rhizomes of some forest herbs and dwarf shrubs of the, 377 Nova Scotia — New Brunswick border region, Variation in selected meristic series in the golden shiner, Note- migonus crysoleucas (Mitchill), in the, 74 Numenius phaeopus, 227 Nyctea scandiaca, 15 Ochrolechia androgyna var. saxorum, 305 oregonensis, 309 Octolasion cyaneum, 395 tyrtaenum, 395 Odocoileus hemionus, 297 virginianus, 32 Oenanthe oenanthe, 317 Oenothera parviflora, 388 Oldsquaw, |, 316 Oligochaeta, Terrestrial, of some New Brunswick caves, with remarks on their ecology, 360 Oliphant, L. W. and S. McTaggart. Prey utilized by urban Merlins, 190 Olynyk, J. and R. Freitag. Collections of spiders beneath snow, 401 Ondatra zibethicus, 96 Onoclea sensibilis, 265, 405 Ontario, central, population of midland painted turtle, Chrysemys picta marginata (Agassiz), Morphological parameters and spring activities in a, 47 Ontario, eastern, Lysurus gardneri, an uncommon stinkhorn observed in, 194 Ontario ecological colloquium, Third annual, 102 Ontario, Fieldfare in, 91 Ontario, Increase in overwintering by the American Gold- finch, Carduelis tristis, in, 165 Ontario, Melanistic Butler’s garter snakes (Thamnophis butleri) at Amherstburg, Ontario, 397 Ontario, New vascular plant records on Pelee and East Sister Islands, Essex County, 384 Ontario, northern, black spruce forest, Changes in small mammal populations after clearcutting of, 41 Ontario, Pteridophytes of the Regional Municipality of Waterloo, 262 Ontario, southern, Distribution of the auricled twayblade orchid (Listera auriculata) in Canada and description of new stations in, 403 Ontario, southern, Some new and intersting grass records from 350 Ontario, southern, Spotted Redshank sighted in, 90 Ontario, southern, Two recent bobcat (Lynx rufus) speci- mens from, 98 Ontario, southwestern, Paspalum ciliatifolium, a grass new to Canada from, 422 Ontario, Swarming of dragonflies noted at Drag Lake, 97 Ontario, The fern Woodsia obtusa (Spreng.) Torrey in, 84 Ontario, Weedy species at terminal grain elevators, Thunder Bay, 158 Opheodrys vernalis, 19 Ophioglossum vulgatum var. pseudopodium, 264 Opuntia compressa, 357 polyacantha, 282 Orchid, Distribution of the auricled twayblade (Listera auriculata) in Canada and description of new stations in southern Ontario, 403 Osmorhiza longistylis, 388 446 Osmunda cinnamomea, 264 claytoniana, 264, 378 regalis var. spectabilis, 264 Ottawa Field-Naturalists’ Club by-laws, The, 114 Ottawa Field-Naturalists’ Club Minutes of the ninety- seventh annual business meeting, 212 Auditor’s report, 215 Balance sheet, 216 Report of council, 213 Statement of profit and loss — CF-N, 217 Statement of profit and loss — O.F.N.C., 218 Ottawa Field-Naturalists’ Club, Notice of change to the by-laws of the, 325 Otter, river (Lutra canadensis), on the north slope of the Brooks Range, Alaska, the, 303 Otter, river, Predation on nesting Glaucous-winged Gulls by, 189 Ovibos moschatus, 78, 81 Owl, Barred, Apparent distraction display by a, 176 Owl, Great Horned, 188 Hawk, 188 Snowy, 15 Oxalis montana, 378 Oxley, D. J. and J. M. Gall. Additional record of the southern flying squirrel from Quebec, 424 Oxley, D.J., R. A. Coutts, and N.G.H. Boyle. Range extensions of the water shrew and mink frog in the James Bay region of Quebec, 322 Oxyria digyna, 14, 82, 368 Oxytropis deflexa var. sericia, 320 Page, G. W., 225 Palomino trout, Genetic variants in Canada of the rainbow trout, Salmo gairdneri, called golden trout and, 93 Panicum agrostoides, 354 capillare, 359 depauperatum, 351 dichotomiflorum, 350, 388 var. geniculatum, 354 Philadelphicum, 388 rigidulum, 350 sphaerocarpon, 350 spretum, 350 virgatum, 357 Pannaria microphylla, 308 Papaver radicatum, 14, 82 Paralysis in a moose calf, Trauma-induced, 94 Parker, G. R. Morphology, reproduction, diet, and behavior of the arctic hare (Lepus arcticus monstrabilis) on Axel Heiberg Island, Northwest Territories, 8 Parker, G. R. The flowering phenology of common vascular plants at Bailey Point, Melville Island, Northwest Territories, 81 Parmelia conspersa, 137 saxatilis, 307 sinuosa, 308 stictica, 308 sulcata, 308 Parmelia cumberlandia, Growth rated in vegetative and reproductive tissue of, after relocation to a new environment, 134 Parmeliella saubinettii, 308 Parnassia montanensis, 368 Parrya arctica, 82 THE CANADIAN FIELD-NATURALIST Vol. 91 Paspalum ciliatifolium, a grass new to Canada from southwestern Ontario, 422 Paspalum ciliatifolium var. muhlenbergii, 423 var. stramineum, 423 Passer domesticus, 187, 191 Passerculus sandwichensis, 191 Paterson, C. G., 74 Peden, A. E. First record of Atka mackerel, Pleurogram- mus monopterygius (Hexagrammidae), in British Co- lumbia, 175 Peden, D. G. Waterfowl use of exotic wild rice habitat in northern Saskatchewan, 286 Pedicularis arctica, 82 groenlandica, 368 hirsuta, 319 spp., 14 sudetica, 82 Pellaea atropurpurea, 264 glabella var. glabella, 264 Pellia epiphylla, 405 Peltigera aphthosa, 307 collina, 308 horizontalis, 308 membranacea, 308 sp., 246 venosa, 308 Pendlebury, G.B. Distribution and abundance of the Prairie Rattlesnake, Crotalus viridis viridis, in Canada, 122 Penny, C. and R. W. Knapton. Record of an American Robin killing a shrew, 393 Penstemon procerus, 368 Perca flavescens, Thermal selection and related behavior in larval yellow perch, 406 Perch, larval yellow (Perca flavescens), Thermal selection and related behavior in, 406 Perch, shiner, in deep water, A Great Blue Heron preying on, 88 Peromyscus leucopus, Movements and habitat use among interacting, as revealed by radiotelemetry, 273 Peromyscus maniculatus, 41, 178, 363 Heinz, Us 185 I<, O7/ Perry, M. S., 97 Perry, T. E., M.S. Perry and J. E. K. Perry. Swarming of dragonflies noted at Drag Lake, Ontario, 97 Pertusaria multipuncta, 309 ophthalmiza, 309 Petasites frigidus, 82 Peterson, R. L., 98 Phacelia purshii, 384 Phalacrocorax auritus, 184 carbo, 3 Phalacrocorax carbo, Le Grand Cormoran en hiver, le long des cétes de la Péninsule de Gaspésie, Québec, 184 Phalaris arundinacea, 388 Phalarope, Northern, 231 Red, 230A Wilson’s, 231 Phalaropus fulicarius, 230 Phasianus colchicus, 32 Pheasant, 32 Phegopteris connectilis, 265 hexagonoptera, 267 OWT Phenacomys intermedius, 41, 96, 239 Phenology of common vascular plants at Bailey Point, Melville Island, Northwest Territories, The flowering, 81 Philohela minor, 141, 227 Phippsia algida, 322 Phoca hispida, 181, 288 Phoebe, Say’s, Breeding status of, in Manitoba, 183 Phragmites communis, 290 Phyllodoce glanduliflora, 367 Physica adscendens, 308 Physostegia virginiana, 388 Pica pica, 297 Picea glauca, 72, 96, 237, 405, 417 mariana, 41, 72, 96, 413, 417 spp. 369 Pinel, H.W. and J.R. Riddell. First record of Anna’s Hummingbird for Alberta, 394 Pintail, 316 Pinus spp., 369 Pipit, Water, 317 Placopsis gelida, 308 Plantago psyllium, 356 Platanthera psycodes, 405 Platismatia glauca, 308 Plectrophenax nivalis, 317 Pleurogrammus monopterygius (Hexagrammidae), First record of Atka mackerel, in British Columbia, 175 Pleuropogon Sabinei, 14, 82, 319 Plover, American Golden, 227 Black-bellied, 227 Piping, 227 Semipalmated, 91, 227, 316 Pluchea purpurascens, 358 Pluvialis dominica, 227 squatarola, 227 Poa arctica, 14 bulbosa, 350 var. vivipara, 356 glauca, 14 spp., 14 Podiceps auritus, 372 Polemonium acutifolium, 322 boreale, 320 Policy, Editorial, 117 Policy, Editor's comments and, 221 Polygonatum pubescens, 378 Polygonum careyi, 355 coccineum, 65 viviparum, 14, 82 Polypodium virginianum, 264 Polystichum acrostichoides, 266, 377 braunii, 267 lochitis, 267 Polytrichum sp., 351 Populus tremuloides, 372, 413 Porcupine, 33, 68, 363 Porina chlorotica, 308 chlorotica var. persicina, 305 Porter, M. J., 91 Porzana carolina, 63 Potamogeton crispus, 368 foliosus, 368 INDEX TO VOLUME 91 447 Potentilla argentea, 388 diversifolia, 368 fruticosa, 246, 368 hyparactica, 82 pulchella, 319 rubricaulis, 319 spp., 367 Vahliana, 82 Powles, P. M., 406 Prairie fires and pronghorn use of cactus, 282 Pratt, P. D., 422 Predators of Snow Goose eggs, Unusual, 317 Prenanthes altissima, 378 trifoliata, 378 Prescott, J., 83 Pringle, J. S., reviews by, 205, 333 Pronghorn use of cactus, Prairie fires and, 282 Prunus sp., 277 Pseudacris triseriata, 20 Pseudocyphellaria anomala, 309 anthrapsis, 307 aurata, 309 crocata, 309 Ptarmigan, 68 Ptarmigan, White-tailed, in southwestern Alberta, Summer habitat use by, 367 Pteridium aquilinum, 378 var. latiusculum, 265 Pteridophytes of the Regional Municipality of Waterloo, Ontario, 262 Pteritis pensylvanica, 378 Puccinellia Andersonii, 12 angustata, 14, 319 nuttalliana, 164 poacea, 14 spp., 14 vaginata, 14 Punpkinseed, 425 Pyrola elliptica, 378 Quebec, Additional record of the southern flying squirrel from, 424 Quebec, Gestation, litter size, and number of litters of the red squirrel (7amiasciurus hudsonicus) in, 83 Québec, Le Grand Cormoran (Phalacrocorax carbo) en hiver, le long, des cétes de la Péninsule de Gaspésie, Québec, 184 Quebec, Range extensions of the water shrew and mink frog in the James Bay region of, 322 Quebec, southern, Distribution and abundance of waterfowl wintering in, | Quebec, southwestern, Aspects of woodcock nocturnal activity in, 141 Quebec, southwestern, Distribution of stream salamanders in, 299 Quedius mesomelinus, 363 Quercus palustris, 388, 423 sp., 277 velutina, 388 Quiscalus quiscula, 65, 187 Racer, blue, 102, 385 Radvanyi, A., 41 Radvanyi, A. An improved design for a small mammal live trap, 399 7 448 Rana septentrionalis, 322 sylvatica, 20, 322 Range extensions, Reporting of, 219 Rangifer tarandus, 96, 317 t. montanus, 312 t. pearyi, 78, 81 Ranunculus eschscholtzii, 320, 368 hyperboreus, 82, 322 nivalis, 82 pallasii, 322 pedatifidus, 14 Sabieni, 82 spp., 14 Rat, bushy-tailed wood (Neotoma cinerea) in Alberta, Range of the, 323 Ratibida pinnata, 389 Rattlesnake, Prairie, Crotalus viridis viridis, in Canada, Distribution and abundance of the, 122 Raven, 32 Recurvirostra americana, 230 Redhead, 248 Redpoll, Common, 317 Redshank, 91 Redshank, Spotted, sighted in southern Ontario, 90 Reed, A. and A. Bourget. Distribution and abundance of waterfowl wintering in southern Quebec, | Referees, Note to authors and, 101 Regulus calendula, 191 Report, Editor’s, 197 Reporting of range extensions, 219 Request for information, colour-banded Semipalmated and Least Sandpipers, 198 Request for information, shorebird color-marking, 198 Request for participants, International Shorebird Surveys, 1977-78, 197 Reynolds, J. W., 360 Reynolds, J. W. Le Ver québécois, (Lumbricus festivus) envahit la Columbie-Britannique, 395 Reznicek, A. A., 350, 384 Rhexia virginica, 355 Rhizocarpon disporum, 308 Rhizomes of some forest herbs and dwarf shrubs of the New Brunswick — Nova Scotia border region, The structure and rate of growth of the, 377 Rhus typhina, 357 Rhynchospora capitellata, 355 Ribes, 414 Rice, exotic wild, habitat Waterfowl use of, 286 Richardson, D. H. S., 305 Riddell, J. R., 394 Riding Mountain National Park, Manitoba, Population fluctuations of wapiti (Cervus elaphus) and moose (Alces alces) in, 1950-1976, 130 in northern Saskatchewan, Riley, J. L., 350 Robarts, D. R., review by, 430 Robin, 191 American, 92 Robin, American, killing a shrew, Record of an, 393 Roe, N. A. and W.J. Stephen. Narwhals (Monodon monoceros) observed near King Christian Island, Northwest Territories, 309 Rorippa islandica, 388 THE CANADIAN FIELD-NATURALIST Vol. 91 Rosa acicularis, 246, 414 palustris, 388 woodsii, 251 Rosa acicularis, Germination requirements of Alaskan, 58 Ross, J., P. M. Powles, and M. Berrill. Thermal selection and related behavior in larval yellow perch (Perca flavescens), 406 i Rounds, R. C. Population fluctuations of wapiti (Cervus elaphus) and moose (Alces alces) in Riding Mountain National Park, Manitoba, 1950-1976, 130 Rubus pubescens, 405 strigosus, 179 Rumex arcticus, 72 maritimus, 388 stenophyllus, 162 Russell, R. H., 77 ; Rutherglen, R. A. and B. Herbison. Movements of nuisance black bears (Ursus americanus) in southeastern British Columbia, 419 Sagittaria sp., 65 Salamander, dusky, 299 small-mouthed, 385 spring, 299 two-lined, 299 Salamanders in southwestern Quebec, stream, 299 Salix, 365 arctica, 8, 82, 368 nivalis, 368 sp., 246 spp., 64, 96, 367, 372 vestita, 368 Salmo aquabonita, 93 Salmo gairdneri, called golden trout and palomino trout, Genetic variants in Canada of the rainbow trout, 93 Salmo trutta Linnaeus X Salvelinus fontinalis (Mitchill), in Alberta, First report of the tiger trout hybrid, 85 Sanderling, 227 Sandpiper, Baird’s, 227 Buff-breasted, 230 Least, 227 Pectoral, 227 Purple, 230, 316 Semipalmated, 227, 316 Solitary, 230 Spotted, 227 Stilt, 230 Upland, 227 Western, 230 White-rumped, 230 Saskatchewan, Nesting and brood ecology of Lesser Scaup at Waterhen Marsh, 248 Saskatchewan, northern, Waterfowl use of exotic wild rice habitat in, 286 Saskatchewan parklands, Horned Grebe breeding habitat in, 372 Saussurea densa, 368 Saxifraga bronchialis, 368 caespitosa, 82 cernua, 82 flagillaris, 82 foliolosa, 322 hirculus, 82 Distribution of 1977 Iyalli, 368 nivalis, 82 oppositifolia, 14, 81, 368 rivularis, 322 spp., 367 tricuspidata, 14, 82 Sayornis saya, 183 Scaup, Lesser, 286 Scaup, Lesser, at Waterhen Marsh, Saskatchewan, Nesting and brood ecology of, 248 “Scavenging” in ecological literature, Usage of the terms “cannibalism” and, 416 Scirpus, 66 spp., 372 Scolochloa festucacea, 251, 372 Scoter, Surf, 3 Scott, D. P. Reporting of range extensions (editorial), 219 Sculpin, slimy, Cottus cognatus, in the Missouri drainage system, Occurrence of the, 415 Scutellaria lateriflora, 405 Sea-bird populations of the north Atlantic, The changing, 102 Seal, ringed, The wolffish, cf. Anarhichos denticulatus, new to the Amundsen Gulf area, Northwest Territories, and a probable prey of the, 288 Seals, ringed and bearded, Disorientation in, 181 Searing, G. F. The function of the bark call of the red squirrel, 187 Sedum rosea, 368 sarmentosum, 388 stenopetalum, 368 telephioides, 384 Selaginella apoda, 263 rupestris, 267 Selection, Thermal, and related behavior in larval yellow perch (Perca flavescens), 406 Senecio congestus, 81 glabellus, 384 lugens, 368 spp., 367 triangularis, 368 Setaria faberi, 350 viridis, 356 Sheep, 32, 87 Sherk, L. C., review by, 430 Shiner, golden, Notemigonus crysoleucas (Mitchill), in the New Brunswick — Nova Scotia border region, Variation in selected meristic series in the, 74 Shorebird color-marking, Request for information, 198 Shorebirds at Long Point, Lake Erie, 1966-1971: seasonal occurrence, habitat preference, and variation in abun- dance, 225 Shrew, arctic, 96 masked, 43, 96, 239 pigmy, 96 short-tailed, 180 smoky, 363 Shrew, Record of an American Robin killing a shrew, 393 Shrew, water, and mink frog in the James Bay region of Quebec, Range extensions of the, 322 Shrubs, dwarf, of the New Brunswick — Nova Scotia border region, The structure and rate of growth of thr rhizomes of some forest herbs and, 377 INDEX TO VOLUME 91 449 Siderits, K., 413 Silene acaulis, 368 Skunk, striped, 33, 71, 248 Skunks, striped, in Delta Marsh, Manitoba, Locations of winter dens utilized by, 289 Smelowskia calycina, 368 Smilacina racemosa, 378 Smilax tamnoides var. hispida, 388 Smith, D. H. The genus Crangonyx (Amphipoda: Gam- maridae) in the central Connecticut River system, 256 Smith, L. C. Editoral Policy (editorial), 117 Smith, L. C. Editor’s comments and policy (editorial), 221 Smith, T.G. The occurrence of a narwhal (Monodon monoceros) in Prince Albert Sound, western Victoria Island, Northwest Territories, 299 Smith, T.G. The wolffish, cf. Anarhichas denticulatus, new to the Amunsden Gulf area, Northwest Territories, and a probable prey of the ringed seal, 288 Smith, T. G. and J. Memogana. Disorientation in ringed and bearded seals, 181 Snake, common garter, 398 island water, 102: smooth green, 19 northern red-bellied, 19 plains hognose, 19 queen, 102 red-sided garter, 19, 70 western plains garter, 19 Snake, garter, Sources of mortality in concentrated popu- lations, 70 Snakes in Manitoba, Life history observations of three species of, 19 Snakes, Melanistic Butler’s garter (Thamnophis butleri), at Amherstburg, Ontario, 397 Snipe, Common, 227 Snow, Collections of spiders beneath, 401 Sobey, D.G. The structure and rate of growth of the rhizomes of some forest herbs and dwarf shrubs of the New Brunswick —Nova Scotia border region, 377 Solanum nigrum, 359 Solidago multiradiata, 368 nemoralis, 351 sempervirens, 357 Somateria mollissima, | m. borealis, 314 Somatochlora williamsoni, 97 Sora at Vermillion, Alberta, Nesting biology of the, 63 Sorex arcticus, 44, 96 cinereus, 43, 96, 239, 393, 400 fumeus, 43, 363 palustris, 322 Sparrow, Chipping, hanged, 69 Sparrow, House, 187, 191, 346 Lincoln’s, 191 Savannah, 191 Song, 191 White-throated, 191 Spartina patens, 350 Spermophilus columbianus, 296 franklinii, 411 lateralis, 297 richardsonii, 410 spp., 70 450 THE CANADIAN FIELD-NATURALIST Sphaerophorus globosus, 308 melanocarpus, 308 Sphagnum, 414 Spiders beneath snow, Collections of, 401 Spilonema revertens, 305 Spiranthes magnicamporum, 384 Spizella passerina, 69 Sporobolus cryptandrus, 351 Spruce, black, 72 White, 72 Spruce, black, forest, Changes insmall mammal populations after clearcutting of northern Ontario, 41 Squirrel, Columbian ground, 296 golden-mantled ground, 297 ground, 70 red, 96, 180, 239 Squirrel, red, Manipulative behavior by a, 417 Squirrel, red, (Tamiasciurus hudsonicus) in Quebec, Gesta- tion, litter size, and number of litter of the, 83 Squirrel, red, The function of the bark call of the, 187 Squirrel, Richardson’s ground, Gestation period and juven- ile age at emergence in, 410 Squirrel, southern flying, from Quebec, Additional record of the, 424 Starling, 187 Steganopus tricolor, 231 Stelfox, J. G. and H. G. Vriend. Prairie fires and pronghorn use of cactus, 282 Stellaria longipes, 82 Stephen, W. J., 309 Stercorarius parasiticus, 316 Sterna paradisaea, 294 Sticta crocata, 307 fuliginosa, 309 limbata, 309 wiegelii, 307 Stinkhorn, Lysurus gardneri, observed in eastern Ontario, an uncommon, 194 Storeria occipitomaculata occipitomaculata, 19 Streptopus roseus, 378 Strix varia, 176 Sturnella neglecta, 191 Sturnus vulgaris, 187 Sugden, L. G. Horned Grebe breeding habitat in Saskatche- wan parklands, 372 Sunfish, green, (Lepomis cyanellus) in the Grand River system, Occurrence of the, 424 Survey, Biological, of the Insects of Canada, Pilot study for a, 199 Surveys, strip aerial, for estimating numbers of wolves on western Queen Elizabeth Islands, Northwest Ter- ritories, Unreliability of, 77 Swallow, Barn, 183 Tree, 191 Symphoricarpos occidentalis, 251 Symposium on wapiti, 325 Synaptomys borealis, 239 cooperi, 41, 180 Syringa vulgaris, 394 Tamias striatus, 180 Tamiasciurus hudsonicus, 96, 180, 187, 239, 417 Tamiasciurus hudsonicus in Quebec, Gestation, litter size, and number of litters of the red squirrel, 83 Vol. 91 Taraxacum hyparcticum, 82 lyratum, 368 pumilum, 82 spp., 14 Teeple, S. M. Reproductive success of Herring Gulls nesting on Brothers Island, Lake Ontario, in 1973, 148 Tern, Arctic, 294 Thalictrum dasycarpum, 384 polygamum, 405 Thamnophis butleri, Melanistic Butler’s garter snakes at Amherstburg, Ontario, 397 Thamnophis radix haydeni, 19 sirtalis parietalis, 19, 70 s. sirtalis, 397 Thank you Earl Godfrey, 198 Thelotrema lepadinum, 309 Thelypteris noveboracensis, 265 palustris, 265 Thomas, J. B. (review by), 201 Thomomys talpoides, 297 Thompson, I. D., reviews by, 103, 332 Thrush, Hermit, 191 Swainson’s, 191 Thuja occidentalis, 405 plicata, 308 Thunder Bay, Ontario, Weedy species at terminal grain elevators, 158 Timm, R. M., 413 Timm, R. M., L.R. Heaney, and D.J. Baird. Natural history of rock voles (Microtus chrotorrhinus) in Minnesota, 177 Titman, R. D., 69 Toad, American, 322 Fowler’s, 102 Toad, boreal, Records of the, from the Yukon and northern British Columbia, 185 Todd, M. C., 312 Townsend, G. F., review by, 328 Tragopogon dubius, 388 Trap, small mammal live, An improved design for a, 399 Tridens flavus, 350 Trientalis borealis, 378 Trillium erectum, 378 undulatum, 378 Tringa erythropus, 90 flavipes, 91, 227 melanoleuca, 227 solitaria, 230 Triodia, 357 Triosteum angustifolium, 389 Triticum sativum, 284 Trout, golden, and palomino trout, Genetic variants in Canada of the rainbow trout, Salmo gairdneri, called, 93 Trout hybrid, tiger, Sa/mo trutta Linnaeus X Salvelinus fontinalis (Mitchill), in Alberta, First report of, 85 Trout, rainbow, Sa/mo gairdneri, called golden trout and palomino trout, 93 Tryngites subruficollis, 230 Tsuga canadensis, 405 Turdus migratorius, 92, 191, 393 pilaris, 91 Turkey, 32 1977 Turnstone, Ruddy, 227 Turtle, midland painted, (Chrysemys picta marginata(Agas- siz), Morphological parameters and spring activities ina central Ontario population of, 47 Turtle, spiny softshell, 102 Tussilago farfara, 388 Twayblade, auricled, orchid (Listera auriculata) in Canada and description of new stations in southern Ontario, Distribution of the, 403 Typha latifolia, 372 sp., 65 Tyrannus tyrannus, 317 Ulmus americana, 85 Uria lomvia, 316 Ursus americanus, 96, 318 Ursus americanus, Movements of nuisance black bears in southeastern British Columbia, 419 Urtica gracilis, 252 User, non-consumptive, The myth of the, 343 Usnea longissima, 308 spp., 307 Vaccinium angustifolium, 178, 414 uliginosum, 246 vitis-idaea, 72 Valerianella intermedia, 389 umbilicata, 389 Valkenburg, P., 303 Vascular plant records on Pelee and East Sister Islands, Essex County, Ontario, New, 384 Vascular plants at Bailey Point Melville Island, Northwest Territories, The flowering phenology of common, 81 Ver québécois (Lumbricus festivus) envahit la Colombie- Britannique, Le, 395 Veronica alpina, 368 Verrucaria maura, 307 Viburnum cassinoides, 405 rafinesquianum, 388 trilobum, 405 Viola sp., 405 Vireo olivaceus, 191 Vireo, Red-eyed, 191 Vole, 291 chestnut-cheeked, 96 heather, 41, 96, 239 meadow, 41, 96, 178, 239 northern red-backed, 96, 239, 417 red-backed, 41 rock, 41 southern red-backed, 178 tundra, 417 Vole, yellow-cheeked, Microtus xanthognathus, in the Northwest Territories, Population dynamics, home ranges, and habitat associations of the, 237 Voles (Microtus xanthognathus), Micro-habitat selection of chestnut-cheeked, 72 Voles, northern red-backed, Clethrionomys rutilus, Inci- dence of the dark color phase in tundra and taiga populations of, 173 Voles, rock, A second population of Microtus chrotor- rhinus in Minnesota with comments on habitat, 413 Voles, rock, (Microtus chrotorrhinus) in Minnesota, Natural history of, 177 Vriend, H. G., 282 INDEX TO VOLUME 91 45] Wanted: data on the seasonal distribution of North Ameri- can Gulls, 325 Wapiti (Cervus elaphus) and moose (Alces alces) in Riding Mountain National Park, Manitoba, 1950-1976, Popu- lation fluctuations of, 130 Wapiti, Symposium on, 325 Warbler, Yellow-rumped, 314 Waterfowl use of exotic wild rice habitat in northern Saskatchewan, 286 Waterfowl wintering in southern Quebec, Distribution and abundance of, | Waterloo, Ontario, Pteridophytes of the Regional Muni- cipality of, 262 Waxwing, Bohemian, 191 Cedar, 191 Weber, W. C., 87 Weedy species at terminal grain elevators, Thunder Bay, Ontario, 158 Weller, W.F. Distribution of stream salamanders in southwestern Quebec, 299 Wells, A. W. Occurrence of the slimy sculpin, Cortus cognatus, in the Missouri drainage system, 415 West, S. D. Manipulative behavior by a red squirrel, 417 Wheater, 317 Whillans, T. H. and E. J. Crossman. Morphological param- eters and spring activities in a central Ontario popula- tion of midland painted turtle, Chrysemys picta mar- ginata (Agassiz), 47 Whimbrel, 227 Whiting, R.E. and P.M. Catling. Distribution of the auricled twyablade orchid (Listera auriculata) in Cana- da and description of new stations in southern Ontario, 403 Wilkes, B., reviews by, 202, 338 Wilkes, B. The myth of the non-consumptive user, 343 Willet, 230 Wishart, R.A. and J.R. Bider. Aspects of woodcock nocturnal activity in southwestern Quebec, 141 Wolf, 15, 96, 323 gray, 77 Wolffia columbiana, 388 punctata, 388 : Wolffish, cf. Anarhichas denticulatus, new to the Amundsen Gulf area, Northwest Territories, and a probable prey of the ringed seal, The, 288 Wolverines, Predation by wolves on, 68 Wolves on western Queen Elizabeth Islands, Northwest Territories, Unreliability of strip aerial surveys for estimating numbers of, 77 Wolves on wolverines, Predation by, 68 Woodcock, American, 141, 227 Woodcock nocturnal activity in southwestern Quebec, Aspects of, 141 Woodpecker, Red-bellied, 385 Woodsia obtusa (Spreng.) Torrey in Ontario, The fern, 84 Woodsia oregana var cathcartiana, 85 Woodwardia virginica, 266 Wright, J., 176 Wrigley, R. E., 28 Xanthium strumarium, 388 Xanthoria candelaria, 308 Xanthoxylum americanum, 85 Xyris difformis, 355 452 Yellowlegs, Greater, 227 Lesser, 91, 227 Yukon and northern British Columbia, Records of the boreal toad from the, 185 Zapus hudsonius, 239 Zapus hudsonius in the Northwest Territories, A recent record of the meadow jumping mouse, 96 THE CANADIAN FIELD-NATURALIST Vol. 91 Zasada, J. C., 58 Zigadenus elegans, 368 glaucus, 388 Zizania aquatica, 286 Zonotrichia albicollis, 191 Zoysia japonica, 350 Index to Book Reviews Botany Brayshaw, T. C. Catkin bearing plants (Amentiferae) of British Columbia, 107 Fassett, N. C. Spring flora of Wisconsin, 334 Ferguson, M. and R. M. Saunders. Canadian wildflowers, 206 Howes, F.N. A dictionary of useful and everyday plants and their comon names, 205 Lindsay, T. S. Plant names, 333 Mohlenbrock, R. H. Guide to the vascular flora of Illinois, 207 Montgomery, F. H. Seeds and fruits of plants of eastern Canada and northeastern United States, 429 Mulligan, G. A. Common weeds of Canada/les mauvaises herbes communes du Canada, 204 Phillips, W. L. and R. L. Stuckey. Index to plant distribu- tion maps in North American periodicals through 1972, 205 Potvin, A. A panorama of Canadian forests, 206 Schwartz, R. Carnivorous plants, 430 Stanton, C. R. Canadian forestry. The view beyond the trees, 335 Vance, F. R., J. R. Jowsey, and J. S. McLean. Wildflowers across the prairies, 334 Viereck, L.A. and E.L. Little. Atlas of United States trees. Volume 2. Alaska trees and common shrubs, 109 Weber, W. A. Rocky Mountain flora, 207 Environment Cameron, T. W. M.andL. W. Billingsley. Energy flow — its biological dimensions. A summary of the IBP in Canada 1964-1974, 336 Golterman, H. L. Physiological limnology — an approach to the physiology of lake systems, 430 Keith, H. F. “Man of the woods”, 335 Kozloff, E. N. Plants and animals of the Pacific North- west, 338 Littlejohn, B. and W. Drew. Superior: The haunted shore, 207 Nettleship, D. N. and P. A. Smith, (eds.). Ecological sites in northern Canada, 110 Ogden, J. G. Il] and M. J. Harvey, (eds.). Environmental change in the Maritimes, I11 Pielou, E. C. Ecological diversity, 336 Ricklefs, R. E. The economy of nature, 208 Scagel, R. F. (ed.). Mankind’s future in the Pacific, 431 Trost, W. R. (chairman). Erosion of land in northwestern Alberta — report and recommendations, 432 Wilson, R. (ed.). The land that never melts: Auyvittuq Natidnal Park, 337 Zoology Bright, D. E. Jr. The insects and arachnids of Canada. Part 2: The bark beetles of Canada and Alaska, 201 Burk, B. Waterfowl studies, 327 Dauphiné, T. C., Jr. Biology of the Kaminuriak population of barren-ground caribou. Part 4: growth, reproduction and energy reserves, 329 Geist, V. and F. Walther, (eds.). The behavior of ungulates in relation to management Gunderson, H. L. Mammology, 203 Harder, W. The anatomy of fishes. Parts I and II, 331 Harrison, H. H. A field guide to birds’ nests, 332 Horn, D. J. Biology of insects, 326 Johnsgard, P. A. North American gamebirds of upland and shoreline, 327 Johnsgard, P. A. Waterfowl of North America, 103 Johnson, W. T. and W. H. Lyon. Insects that feed on trees and shrubs: an illustrated practical guide, 105 Lumi, M. Wolf... kill. The wilderness called Shunka, 331 Massé, G. et J.-R. Mongeau. Répartition géographique des poissons, leur abondance relative et bathymeéetrie de la région du Lac Saint-Pierre, 330 May, C. P. A book of Canadian mammals, 202 Menzies, J. I. Handbook of common New Guinea frogs, 103 Miller, D. R. Biology of the Kaminuriak population of barren-ground caribou. Part 3, 201 Miller, F. L. Biology of the Kaminuriak population of barren-ground caribou. Part 2, 200 Mongeau, J.-R., A. Courtemanche, G. Massé, et B. Vicent. Cartes de répartition géographique des espéces de poissons au sud du Québec, d’aprés les inventaires icythyologiques effecués de 1963 a 1972, 330 Mongeau, J.-R. Méthodes de péche expérimentale, en eau douce, a usage du biologiste et du technicien de la faune, 330 Mongeau, J.-R. et G. Massé. Les poissons de la région de Montreal, la péche sportive et commerciale, les en- semencements, les frayéres, la contamination par le mercure et les PCB, 330 Nail, F. America’s master of bee culture. The life of L. L. Langstroth, 328 Neill, W. T. Reptiles and amphibians in the service of man, 427 Orr, R. T. Vertebrate biology, 429 Palmer, R. S. (ed.). Handbook of North American birds. Volumes 2 and 3, Waterfowl, Parts | and 2, 104 Parker, G. W. An investigation of caribou range on Southampton Island, N.W.T., 203 Rue, L. L., Ill. Pictorial guide to the mammals of North 1977 INDEX TO VOLUME 91 453 America, 333 Walker, E. P. Mammals of the world, 107 Serle, W. and G. J. Morel. A field guide to the birds of West Africa, 428 Other Books Struhsaker, T. T. The red colobus monkey, 427 Stanley, D. J. and D. J. P. Swift (eds.). Marine sediment Trefethen, J. B. An American crusade for wildlife, 328 transport and environmental management, 209 Instructions to Contributors Content The Canadian Field- Naturalist is a medium for publica- tion of original scientific research papers in all fields of natural history that have relevance to Canada. As the journal has a flexible publication policy, items not covered in the traditional sections (Articles, Notes, Letters, News and Comment, and Book Reviews) can be given a special place provided they are judged suitable. Naturalists are also encouraged to support local natural history publications. Manuscripts Please submit, in either English or French, three complete manuscripts written in the journal style. The research reported should be original. 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Almost all manuscripts accepted for publication have undergone revision—sometimes extensive revision and reappraisal. The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain the scientific quality and overall high standards of the journal. TABLE OF CONTENTS (Concluded) Notes (continued) Movements of nuisance black bears (Ursus americanus) in southwestern British Columbia R. A. RUTHERGLEN and B. HERBISON 419 Paspalum ciliatifolium, a grass new to Canada from southwestern Ontario WILLIAM J. CRINS, PAUL D. PRATT, and DANIEL F. BRUNTON 422 Additional record of the Southern Flying Squirrel from Quebec D. J. OXLEY and J. M. GALL 424 Occurrence of the Green Sunfish (Lepomis cyanellus) in the Grand River System EDWARD KoTT and GREGORY B. HUMPHREYS 424 Book Reviews Zoology: The Red Colobus Monkey — Reptiles and amphibians in the service of man— A field guide to 427 the birds of West Africa — Vertebrate biology. Botany: Seeds and fruits of plants of eastern Canada and northeastern United States — Carnivorous 429 plants Environment: Physiological limnology - an approach to the physiology of lake ecosystems — 430 Mankind’s future in the Pacific — Erosion of land in northwestern Alberta—report and recommendations. New Titles 433 Index to Volume 91 Compiled by R. EMERSON WHITING 436 Mailing date of previous issue 3 November 1977 SPECIAL THANKS The Publications Committee of The Ottawa Field-Naturalists’ Club acknowledges with special thanks the contributions of the National Research Council of Canada and The Canadian National Sportsmen’s Show toward the publication of this volume. THE CANADIAN FIELD-NATURALIST Volume 91, Number 41977 Viewpoint The myth of the non-consumptive user BRIAN WILKES 343 Articles Some new and interesting grass records from southern Ontario P. M. CATLING, A. A. REZNICEK, and J. L. RILEY 350 Terrestrial Oligochaeta of some New Brunswick caves with remarks on their ecology DONALD F. MCALPINE and JOHN W. REYNOLDS 360 Summer habitat use by White-tailed Ptarmigan in southwestern Alberta PATRICK W. HERZOG 367 Horned Grebe breeding habitat in Saskatchewan parklands LAWSON G. SUGDEN B72 The structure and rate of growth of the rhizomes of some forest herbs and dwarf shrubs of the New Brunswick — Nova Scotia border region DOUGLAS G. SOBEY and PETER BARKHOUSE 377 New vascular plant records on Pelee and East Sister Islands, Essex County, Ontario CRAIG A. CAMPBELL and A. A. REZNICEK 384 Notes Geographic variation in Dunlins, Calidris alpina, of North America M. RALPH BROWNING 391 Record of an American Robin killing a shrew CHERYL PENNY and RICHARD W. KNAPTON 393 First record of Anna’s Hummingbird for Alberta HAROLD W. PINEL and JOHN R. RIDDELL 394 Le ver québécois (Lumbricus festivus) envahit la Colombie-Britannique JOHN WARREN REYNOLDS 395 Melanistic Butler’s Garter Snakes (Thamnophis butleri) at Amherstburg, Ontario P. M. CATLING and W. FREEDMAN 397 An improved design for a small mammal live trap ANDREW RADVANYI 399 Collections of spiders beneath snow J. OLYNYK and R. FREITAG 401 Northern range extension for the Brassy Minnow in northeastern Alberta DAVID K. BERRY 402 Distribution of the auricled twayblade orchid (Listera auriculata) in Canada and description of new stations in southern Ontario R. EMERSON WHITING and PAUL M. CATLING 403 Thermal selection and related behavior in larval yellow perch (Perca flavescens) JAY Ross, P. M. POWLES, and MICHAEL BERRILL 406 Gestation period and juvenile age at emergence in Richardson’s Ground Squirrel GAIL R. MICHENER 410 A second population of rock voles, Microtus chrotorrhinus, in Minnesota with comments on habitat RICHARD R. BUECH, ROBERT M. TIMM, and KARL SIDERITS 413 Occurrence of the Slimy Sculpin, Cottus cognatus, in the Missouri drainage system ALAN W. WELLS 415 Usage of the terms “cannibalism” and “scavenging” in ecological literature MARTIN K. MCNICHOLL 416 Manipulative behavior by a Red Squirrel STEPHEN D. WEST 417 Harassment of an elk calf by bison BRIAN R. MAHAN 418 ISSN 0008-3550 concluded on inside back cover ne ie wi os al Wis uh w ce at aye Sta A: CIme Bookbinding Co., Inc. 300 Summer Street Beston, Mass. 02210 TL zeal eRe - ‘ cae EP gs eo nae ; RP Pca D LEN Se Cte Tn nee ae ory eee nen ret ee AF ’ Lanett re wee oO pe root ey eter ne ern y : , - ~ prs “ty AP Te PPI OS 7 otal