24 A atte tect tat otek i 8 Ban lekened 25o Ee 33 reed cP Soe e hina hy ted ri Sed Ae tase Ant hae Sarai. . hse tact See ak S59 ees cts 3 -t2H eR Note =. pee ae ars ST ae a wt Dey san ee Nereus tates preowes Me Rta Oe hh ei be Be Chk a ta cake “ePe rains ores SOPH Ph. we pos Eh Eme™ bhi ie BT aie tes bs fon tl te ies hth ie oe Vg eS eS . a TN ete sin fat en lon Sa Seeks: toe deh te VA eas BUNS “te Ae ——4 meme Nete tesesens Pte eres Qre ee bs ta ho be aah Oa TI aR SP. tet POTTY Ny yt abe AEE NS PN BOT b abe Sie keke de fa Soke ws *8 tat 2 en (\ The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 116, Number 1 \ } January-March 2002 a SEMAN TED a eee, The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Her Excellency The Right Honourable Adrienne Clarkson, C.C., C.M.M., C.D. Governor General of Canada His Excellency John Ralston Saul, C.C. 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 infor- mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain- ing or restoring environments of high quality for living things. Honorary Members Edward L. Bousfield Bruce Di Labio George F. Ledingham Robert W. Nero Donald M. Britton R. Yorke Edwards John A. Livingston E. Franklin Pope Irwin M. Brodo Anthony J. Erskine Stewart D. MacDonald William O. Pruitt, Jr. William J. Cody John M. Gillett Hue N. MacKenzie Joyce and Allan Reddoch Francis R. Cook W. Earl Godfrey Theodore Mosquin Mary E. Stuart Ellaine Dickson C. Stuart Houston Eugene G. Munroe Sheila Thomson 2002 Council President: Eleanor Zurbrigg Ronald E. Bedford Francis R. Cook David W. Moore . . J Rosanne Bishop Barbara Gaertner Robert Roach Bie eeeiments:. Rey: Jol Irwin Brodo Diane Lepage Stanley Rosenbaum Gary McNulty John Cameron Diane Holmes Louise Schwartz Recording Secretary: Ken Allison William J. Cody David Hobden David Smythe Kathy Conlan Beverly McBride Dorothy Whyte Treasurer: Frank Pope To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, Box P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net. For information on Club activities telephone (613) 722-3050 or check http//www.achilles.net/ofnc/index.htm The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP), toward our mailing cost. PAP Registration number 09477. Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1RO; (613) 269-3211; e-mail: feook @achilles.net Copy Editor: Wanda J. Cook Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 759-1374 Book Review Editor: Dr. J. Wilson Eedy, 9074-32 Side Road, R.R. 1, Glencairn, Ontario LOM 1KO; e-mail: terfa@ geoniger.com; edithheedy @ prepaidlegal.com Associate Editors: Robert R. Anderson Robert R. Campbell Brian W. Coad W. Earl Godfrey Charles D. Bird Paul M. Catling Anthony J. Erskine William O. Pruitt, Jr. Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication should be addressed to the Editor and sent by postal mail (no courier, no post requiring signature on delivery). Exception: book reviews should go directly to Book Review Editor. Subscriptions and Membership Subscription rates for individuals are $28 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $28 (individual) $30 (family) $50 (sustaining) and $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and mem- bers (including USA) must add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at $28 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications Mail Agreement number 40012317 PAP Registration number 09477. Return Postage Guaranteed. Date of this issue: January—March 2002. Cover: Wood Bison, Bison, bison bison photographed at Wood Buffalo National Park. Photo courtesy W.A. Fuller. See: Canada and the “buffalo” pages 141-159. THE CANADIAN FIELD-NATURALIST Volume 116 2002 THE OTTAWA FIELD-NATURALISTS’ CLUB OTTAWA CANADA ee i ee 4c i t i! ty i ; \ 2; 4 we - AN, vi = a | yh ; Mie 4 ‘ ciated ve © : t ' mh ’ £ he? Ll AMAT wet oy. i0s rae oe | ell oe ii ay Ric? bie eoivih (ollwomnedit yen SBM NCR il EIN : Sans a pe : 1 tr tol hale. a eek sO 4 ag eng 9 a bering! Wy oF Delt © ie, ‘ ¥ 2 7 Saft : Wed o ety - i ev, 7 Vy i os ra ak Wis ie Cae eS ae s jeneuell, vi ? P J & — ® bo? te a y Peo Ras. ae wd Chetek WOE ’ La af aN? op ; hea Le eth) 3 MPRANY ee At Rei ‘i es) 4 bing, fw) Ser a H hy ew Op een heal a, Te ON F hh Phin te ae’ Cana Rr ti Capi 8 6) agg Me Agate B94 eels ain vf Pad ial, Pills) wale Sanpete: < AM cree homers Aare I 4 w Py re a 2 m sl i) Ay ptecvepaie at # Aww sci ~ Peeh. Hw rs , @y de ce x “9 The Canadian Field-Naturalist Volume 116, Number 1 January—March 2002 Spread and Disappearance of the Greater Prairie-Chicken, Tympanuchus cupido, on the Canadian Prairies and adjacent areas C. STUART Houston! 1863 University Drive, Saskatoon, Saskatchewan S7N 0J8 Canada Houston, C. Stuart. 2002. Spread and disappearance of the Greater Prairie-Chicken, Tympanuchus cupido, on the Canadian prairies and adjacent areas. Canadian Field-Naturalist 116(1): 1-21. In western Canada, the Greater Prairie-Chicken (Tympanuchus cupido) has come and gone. Following closely on agricul- tural settlement, thriving on grain as an alternate food, the “Pinnated Grouse” reached Winnipeg in 1881 and Carberry, Manitoba, in 1886; Indian Head, Saskatchewan, in 1895; and Hanna, Alberta, in 1911. It spread as far northwest as Lac la Biche, Alberta. Once half the land in a given area was broken, this species diminished in numbers, retreating to local areas of thick grass around sloughs and lakes. By the late 1930s almost all were gone. Habitat factors such as fragmentation and separation of grasslands were further accentuated by cattle overgrazing, burning, and drought, and then by hybridization of surviving, isolated birds. By accessing records for the prairie provinces from nine unpublished or not-readily-available sources (unavoidably skewed by a preponderance of Saskatchewan records), I offer documentation of increases and decreases, as well as an assessment of the species’ status in northeastern Montana and northwestern Ontario. I have also catalogued 131 Northern Great Plains egg sets from 36 North American oology collections. Together this material allows a more complete assessment than has been possible previously. Key Words: Greater Prairie-Chicken, Tympanuchus cupido, spread, decline, extirpation, ecologic factors. On the Canadian grasslands, the Greater Prairie- Chicken (Tympanuchus cupido) has come and gone. Because there is concern for many other species of grassland birds, including the Sharp-tailed Grouse (Tympanuchus phasianellus), and because two other species, the Sage Grouse (Centrocercus urophasi- anus) and Burrowing Owl (Athene cunicularia), may be facing extirpation in Saskatchewan, it seems worthwhile to summarize the spread and demise of the Greater Prairie-Chicken in western Canada and adjacent North Dakota. Are there common threads or lessons to be learned? In this review I bring together, before it is too late, published and previously unpublished information. To provide localities and dates, and especially breed- ing records, I made particular use of: (a) the Andrew Graham/Thomas Hutchins combined report from Hudson Bay, 1772, (b) David Douglas’ Manitoba observation of 1827; (c) responses to Mrs. Isabel Priestly’s request (1943) for Saskatchewan records; (d) unpublished responses to my request for addi- tional Saskatchewan records in 1956; (e) George J. Mitchell’s 1959 Alberta’s Upland Game Bird Resource (1959); (f) Margaret Belcher’s 1961 report of museum records in Saskatchewan; (g) Dale Hjertaas’ 1988 summary of Saskatchewan records since 1966; (h) Anita Steeg Kovacs’ 1984 summary of newspaper accounts in southern Manitoba, 1884-1949, (1) a Montana sportsman’s magazine article that adds extreme northwest Montana to their maximum range, (j) Harry Lumsden’s previously unpublished reports that add northwestern Ontario to their range, and (k) oology records for the three prairie provinces and the Dakotas in major museums. This information has been brought together in one place for use by future researchers, even though most of the Saskatchewan locations have been mapped in Smith (1996). The dearth of readily avail- able accounts largely explains the paucity of western Canadian information in Schroeder and Robb (1993) Problem of name The names used for prairie grouse could hardly be more confusing. Sadly, “Prairie Chicken” was, and still is, the name almost universally applied to the Sharp-tailed Grouse through western Canada. Ernest E. Thompson [Seton] in his 1890 account of the Birds of Manitoba, used “Prairie Chicken” for his account of the Sharp-tailed Grouse, and used “Prairie Hen” or Pinnated Grouse for what was then known to ornithologists south of the boundary as the Greater Prairie-Chicken. Seton’s usage followed that of Macoun (1883). Macoun and Seton thus elevated the use of “Prairie Chicken” for the Sharp-tailed Grouse far beyond that of a simple and misleading folk ~~ were ~~. ee eee Zz THE CANADIAN FIELD-NATURALIST name and gave such use scientific standing for more than half a century. The general public and even game guardians and museum employees used “Prairie Chicken” much more often for the Sharp- tailed Grouse than for the Greater Prairie-Chicken. For this reason, most popular accounts that refer to “prairie chickens” are suspect. In fact, nearly all of them refer to the Sharp-tailed Grouse. Johnston and Smoliak’s helpful review of the Greater Prairie- Chicken in 1976 fell into this trap when they gave credence to Bulyea (1901) who unquestionably referred to Sharptails between Moosomin and Qu’ Appelle, Saskatchewan; they might better have quoted Rutherford (1914), who did recognize the presence of Greater Prairie-Chicken, and, differenti- ated between the two species. To avoid confusion, Cooke (1885), Roberts (1936) and Johnsgard (1973) used the term “Pinnated Grouse.” Although “Pinnated Grouse” referred to only one subspecies of the Greater Prairie-Chicken, and although: common names are no longer assigned to subspecies, “Pinnated” is the only term that unequivocally indi- cates that past observers, at a given time or place, saw the Greater Prairie-Chicken. Problem of identification In an ideal world, one would anticipate few prob- lems of mistaken identification between two grouse species that show such marked differences, one from the other. First, those reports that specifically men- tioned the dark, rounded or squared tail, and the heavily barred underparts, offer proof of the Greater Prairie-Chicken, in sharp contradistinction to the light-coloured, elongated ‘pin-tail’ and finer v- shaped breast markings of the Sharp-tailed Grouse. Although the Ruffed Grouse (Bonasa umbellus) also has a rounded tail, its restriction to wooded areas should exclude it from consideration in most plains localities. Second, the mating rituals at the “booming grounds” of the Greater Prairie-Chicken and the “dancing grounds” of the Sharp-tailed Grouse are different. Third, as a game bird, individuals from each fall kill were available for close inspection, pro- viding in-the-hand identification opportunities not available for many non-game species. The most credible observers are those who mentioned the two species together, for these folks clearly had an opportunity for comparison. Nevertheless, one would be naive to expect uniform accuracy, because wishful thinking, a fertile imagination, or a faulty memory may on occasion have contributed to occa- sional and innocent misidentifications, particularly of sightings since 1940 — and we know not which or how many of these have been reproduced here. Initial range It is somewhat ironic that the Greater Prairie- Chicken was becoming extirpated in the eastern Vol. 116 - United States even as it spread west across the plains. For example, it disappeared from Tennessee in 1850, Kentucky in 1874, Arkansas in 1913, and Ohio in 1934 (Schroeder and Robb 1993), and its decline in northern Illinois began as early as 1850 (Schorger 1944). “At one time it was the leading upland game bird of the grasslands of central North America” (Robel et al. 1970). In Wisconsin in 1850, “Two sportsmen, with one dog, generally bag from fifty to eighty in a day” (Hoy 1852). “Thousands of barrels of them” were sold by market hunters (Rue and Allen 1973). At the time of the return trip of the Lewis and Clark expedition in 1806, Greater Prairie-Chickens were present in Illinois. The sighting “highest up” the Missouri was at the James River, (now Yankton, South Dakota), on 2 September 1806 (Burroughs 1961). T.S. Roberts (1936) stated “There were me Pinnated Grouse in Minnesota in the days of the early explorers. ... [this species] extended its range westward and northwestward just as the country was settled, keeping pace with the forefront of grain- fields.” Through 1850 to perhaps 1880, the range of overlap between the Pinnated Grouse and the Sharp- tailed Grouse range was very narrow, because the one species ceased where the other began (Johnsgard and Wood 1968). The 18"-century record for Ontario Previous accounts have overlooked Pennant’s 1792 statement, “Is frequent about a hundred miles up Albany river, in Hudson’s Bay.” This statement is apparently derived from the Andrew Graham unpub- lished manuscript E.2/9, Hudson’s Bay Archives, Provincial Archives of Manitoba, written probably in 1772: “Pinnated Grous [sic] is found about Henley Settlement in Hudson’s Bay. Legs covered with soft brown feathers, toes naked and pectinated. The tufts which distinguish this species from all others are rooted high on the neck, not far from the hindpart of the head.” (Houston et al. 2002). Henley House was situated just below the junction with the Kenogami River, about 160 km upstream from Albany on Hudson’s Bay. In 1828, the farthest north known record in Ontario of the Greater Prairie-Chicken was in Essex County, about 1000 km south of Henley House. As settlers cleared land, it spread eastward to Toronto by 1858 and north to Holland Marsh at the south end of Lake Simcoe by 1875. It then retreated, and was extirpated from the province by 1924 (Lumsden 1966). Next, the species spread north in Michigan to Sault-Ste. Marie, then gradually colonized Manitoulin Island, from west to east, between 1941 and 1962, before being swamped through hybridiza- tion with the Sharp-tailed Grouse, newly arrived from the north (Peck and James 1983). 2002 HOUSTON: GREATER PRAIRIE-CHICKEN ON THE CANADIAN PRAIRIES 3 Overlooked record near present Winnipeg All previous accounts, save that of Roberts (1936), have overlooked an important record. In the first six days of August 1827, David Douglas, under the heading Tetrao cupido, “killed several birds of this species between Red River and Pembina in 49° north latitude. This may, perhaps, be found to be its most northern range” (Douglas 1829). Douglas arrived at Fort Garry, within present Winnipeg, on 12 July 1827 and stayed until 10 August. He made a number of one-day botanizing trips and one two-day trip on 23-24 July from Fort Garry to Whitehorse Plain, 29 km west up the Assiniboine River. On 1 August he went on horseback to a low hill about 27 km east of Fort Garry (Douglas 1914). His Greater Prairie-Chickens must have been collected much closer to Fort Garry than to Pembina. In view of Douglas’ single record of this species within Manitoba in 1827, Roberts’ claim that there were none in Minnesota through 1850 is probably an error. Spread into Northwestern Ontario (Figure 1) The only published observation in northwestern Ontario, of “occurrence” with “no indication of breeding,” is in an eight-page mimeographed list, Birds of the Canadian lakehead area, by K. Denis (1940, cited by Peck and James 1983: 137). Nevertheless, Greater Prairie-Chickens were present between 1920 and 1940 near Warroad, Minnesota, at the southwestern corner of Lake of the Woods, and east along the Rainy River, the United States— Canada boundary, to Fort Frances (The last booming was heard 5 km southeast of Warroad in 1963; J. L. Ruos, Leader, Grouse Research Project, Minnesota Game and Fish, letter to Harry G. Lumsden, 16 December 1963). It also occurred within northern Minnesota as far east as Duluth until 1952 (Green and Janssen 1975). Slightly farther north, near Dryden, Ontario, Chief Ranger John Anderson in 1944 reported to Lumsden that he saw 10 to 15 grouse with yellow-orange neck patches displaying near the four-mile corner of the Rice Lake loop road, east of Quibell. Ontario Conservation Officer A. R. Olsen (six-page report to Lumsden, 1959) inter- viewed long-time farmers west of Dryden and deter- mined that, after they cleared the bush for farming, Sharp-tailed Grouse moved in and increased until the 1920s, accompanied by a few Greater Prairie- Chicken, which were preferred for food because of their larger size. On 8 April 1959, 5 km north and | km east of Minnitaki post office west of Dryden, Olsen observed a dancing ground of 2 hectares on a stubble field with six birds. When flushed, two were positively identified as Pinnated Grouse and the other four as Sharptails. The above observations appear to add a new species to at least the hypotheti- cal bird list for Northwestern Ontario. Spread into Manitoba (Figure 1) The spread of the Greater Prairie-Chicken into, and across, Minnesota, beginning about 1850, has been well documented by Roberts (1936). Cooke (1885) reported from Moorhead, Minnesota, its inva- sion of the Red River valley of Manitoba: “Its north- ern range has always fallen short of our northern boundary until last summer, when they invaded Manitoba. Mr. C. W. Nash of Portage la Prairie, says: ‘This autumn we had a curious influx of the Pinnated Grouse. I imagine that they, like some other birds, are following up civilization, for until last year (1884) none were seen by the Indians and half-breed hunters.’” The Greater Prairie-Chicken then spread west roughly in concert with white settlement. In Manitoba it reached Winnipeg in 1881, Portage la Prairie in 1882 (Nash in Cooke 1888), Carberry in 1886 (Thompson 1890), and Aweme/Treesbank in 1888 (Criddle 1929). Alex McArthur gave a talk on winter birds in January 1887 and reported that the previously absent Pinnated Grouse was “now quite common ... The reason for its rather sudden appear- ance and rapid increase seems to be that it has fol- lowed the wheat fields.” C. P. Forge collected at least 17 Greater Prairie-Chicken egg sets near Carman, Manitoba, 1900-1904 (Table 1, Houston and Bechard 1987b). Four decades after their arrival in Manitoba, they were still at a booming ground near Carberry (Symons 1967). They spread north as far as Swan River (specimen in 1938; Minish 1987a), Peonan Point (specimen) and Grand Rapids (sight record) (Godfrey 1966). According to T. Schindler, they reached their highest numbers at Oak Hammock Marsh in the 1920s (Gardner 1981). Spread into Saskatchewan (Figure 2) In Saskatchewan, Greater Prairie-Chickens spread and multiplied rapidly. One was killed at Indian Head late in 1895 (Macoun 1900). The first Saskatchewan nest was found at Crescent Lake, near Saltcoats, in 1897 (Table 1), where from 1910 to 1920 they were as common as the Sharp-tailed Grouse (Baines 1956), a claim made also by Ferry (1910) at the Quill Lakes in 1909. Greater Prairie- Chickens reached Tullis after 1906 (Roy 1996). Their presence on intensively farmed land near Regina was short-lived, but they were numerous 5 km west of that city on 30 September 1913 (H. H. Mitchell, in Belcher 1961), and were also seen 8 km south of that city on 27 April 1913 (Buchanan 1914). They reached the Battleford district in 1913, and there were “large flocks” at Lloydminster in 1914 (Bradshaw 1915). In 1914, J. D. Soper (1970) found “a few” at a ranch 10 km south of Broadview, but when he returned from late April to early June 1921, he saw none, though residents reported a few still —_ eee ee eee SASKATCHEWAN li ts of ee ef OV ND ulvihill Kalevala Birtle Langruth Beulah Bie Delta Beach c= MacDonald, eOakland Douglas e Brandon | | \—reesbank Broomhi ( roomhill we Boissevain, Roland® * *Pilot Mound Portage la “Carberry _/ Prairie Wawanesa THE CANADIAN FIELD-NATURALIST e ak Stonewall Vol. 116 ps a OlNWLNO Quibell Dryden 49°N 6 __ Frances Ss p ae = ae el 94°W "MINNESOTA~ Warroad Rares Rainy*—~_ River FIGURE |. Localities mentioned in Manitoba. present. In 1915 George Lang reported large flocks of prairie grouse at Indian Head, approximately one- half being Greater Prairie-Chicken or hybrids (the only mention of hybridization located in the Saskatchewan game reports). At Indian Head in the winter of 1916-1917 they were sufficiently common that George Lang, voluntary game guardian, observed seven dead birds within one mile; they had been killed by flying into telephone wires (Bradshaw 1917). They appeared at grain baits at Raymore in the winter of 1916-17 with flocks of Sharp-tailed Grouse (Charles Harris in Wayne C. Harris, unpub- 2002 JOnion Lake oe Maidstone, BZ CA | Cut Knife, ALBERTA ol ND ° —— ee ee — —— s e South Leader | | | o°| wv YPRESS Cypress Hills Prov. P. HILLS ark Altawan e Govenlock lished manuscripts). In 1918, H. F. Ward at Pennock near Saltcoats reported a “surprising number” of Pinnated Grouse, to six or seven in other flocks, and several singles (Bradshaw 1919), HOUSTON: GREATER PRAIRIE-CHICKEN ON THE CANADIAN PRAIRIES R lve Ya eonipe Lake Lac Ile-a- la-Crosse \ i Montreal Lake "sal Lake e Snowden River / Edam Henribowg, Prince Albert S 5s Nipawin Codette Davis? cart 6. S .Battleford y c el isdale Sy ) & Radisson, B arden Q Ss Kelvington e,.. BS Lintlaw Quill Lakes Saskatoon eOundurn Watrous ALLAN HILLS Nokomis, Simpson, Bladworth, BN ge Davidson. we Dafoe Sheho e & NY gS Kamsack eRaymore evorkton Ituna + Saltcoats| e e Mountain Tullis, Lake ' Melville, Holdfast Qu’Appelia Katepwae Qu’Appelle, , ¥ ee? REGINA ‘ss gio *Abernethy Saskatchew?? River Broadview® : en ; 2 Whitewood Moosomine Craven Mortlach e Moose Jaw Old Wives Lake ° | soe Langbank \ - carlyle Wauchope Arcola. "Re dvers Storthoaks, Frenchma, Gainsborough Wood Mountain Willow Bunch gL e Estevan Val Marie Coronach Killdeer, North Portal | NORTH DAKOTA | 104° FIGURE 2. Localities mentioned in Saskatchewan. At Ituna they were first seen about 1910, five or six years after initial settlement, when little land was yet broken; H. M. Rayner saw an occasional covey until the fall of 1943 when he flushed eight birds 3 km NNE of Ituna (letter, 4 March 1956). Fred G. including one covey of eleven, up 6 THE CANADIAN FIELD-NATURALIST oO &—__BRINSH CoLumBia a Watertd n Park. WASHINGTON ~T IDAHO | cies = Ak | BME Rabe A Vol. 116 Lesser Slave ways ae . : | Primrose Ss Lac la Biche Cold Lake Saskatepe, | 54° a4 river 3) | Beaverhill [E Camrose ° — Bawif *Daysland ——_— NHL SVS a m \_ Cardston ¢ e JENS *Mountain View __, Waterton-Glacier\::: i International Peace . 16° P. FIGURE 3. Localities mentioned in Alberta. Bard found them still common, evidently breeding, at Dafoe in both 1928 and 1929, and collected an adult specimen on 25 April in the latter year (Belcher 1961). The Greater Prairie-Chicken reached the northern limit of its range in longitude 102° at Kamsack in 1907 (Fraser 1961). Farther west it reached Edam in 1913 where W.E. Lake collected one on 13 September (Belcher 1961). By 1914 it had reached the Onion Lake Indian Reserve, near the Alberta boundary north of Lloydminster (Bradshaw 1915). The species was already present at Somme when Wallace Black homesteaded in 1920, and it reached Tisdale in 1923 (Houston and Street 1959). By 1924 2002 HOUSTON: GREATER PRAIRIE-CHICKEN ON THE CANADIAN PRAIRIES 7 TABLE 1. Greater Prairie-Chicken egg sets, Northern Great Plains (see acknowledgments for institute initials) # sets sets eggs/set c eggs Locality Day/Mo Year # eggs Collector Institution Unknown 4 3 10.67 32 unknown — = 1905 7 J Kowal PMA unknown — = — 12 J Kowal PMA unknown == — (1) J Kowal PMA unknown a 20 May 1930 13 oa ASC Alberta (AB) 2 2 11.50 23 Stony Plain — 1901 14 SSS Stansell? PMA Heatherdown — — 9 M Pollack/Wilby PMA Saskatchewan (SK) 13 8 10.88 87 Crescent L 24 May 1897 11 F Baines WFVZ Saltcoats — 1915 — N Jowsey PMA Saltcoats — 1915 (2) N Jowsey PMA Melville 30 May 1920 10 HC Grose SMNH Wauchope 20 May 1923 (4) HH Pittman USask Wauchope 10 May 1924 (1) HH Pittman SMNH North Portal 3 May 1925 12 GF Abbey PMYU S of Gainsborough 12 May 1925 14 GF Abbey RBWWEF Wauchope 25 May 1925 (1) HH Pittman USask 5 mi nw Antler ND* 6 June 1925 1] GL Davy WFVZ 5 mi nw Antler ND* 29 May 1927 9 GA Withey WFVZ 4 mi nw Antler ND* 8 June 1927 10 GL Davy WFVZ 5 mi nw Antler ND* 14 June 1927 10 GA Withey SBCM Note: those four with ND* probably were in Saskatchewan Manitoba (MB) 32 30 10.03 301 Shoal Lake 8 June 1894 6 ne E Arnold WFVZ nr Reaburn 9 June 1894 12 W Raine NMC Morris 17 May 1896 u PB Peabody MPM Beulah 29 Apr 1899 2 ne AJ Dennis DMW Long L nr Reaburn 10 June 1899 8 W Raine WFVZ Carman 30 May 1899 13 JW Preston ROM Carman — — 8 JW Preston OSUM nr Roland 25 May 1900 8 CP Forge WFVZ Carman 28 May 1901 9 CP Forge OSUM Barnsley 29 May 1901 13 WC Bradbury DM Salterville 30 May 1901 8 CP Forge NYSM Salterville 10 June 1901 1] CP Forge WFVZ Myrtle 15 May 1902 9 ne CP Forge WFVZ Salterville 17 May 1902 8 CP Forge WFVZ Salterville 17 May 1902 11 CP Forge WFVZ Myrtle 19 May 1902 8 CP Forge MVZ nr Roland 31 May 1902 10 CP Forge WFVZ Myrtle 31 May 1902 17 CP Forge FMG Myrtle 21 June 1902 10 CP Forge WFVZ Myrtle 21 June 1902 15 CP Forge WFVZ Madford/Wadhope? 2 July 1902 13 J Labarthe UNR Myrtle 5 June 1903 10 ne GE Congdon WFVZ Elm Point 20 June 1903 10 CP Forge WFVZ e of Carman 24 May 1904 14 CP Forge AMNH Salterville 24 May 1904 9 CP Forge USNM MacDonald 22 June 1905 6 CP Forge?/Wilby PMA Lake Winnipeg 2 June 1905 10 DWilby NMC Douglas 1 June 1916 eae! CH Young NMC Kalevala 28 May 1921 12 ES Norman SBCM Kalevala 30 May 1921 13 ES Norman UMMZ Oaklands 11 June 1927 6 AG Lawrence NMC St. Francis Xavier 4 June 1928 12 AG Lawrence NMC Continued —_ woe eee ee ee 8 THE CANADIAN FIELD-NATURALIST TABLE 1. Continued # sets sets c eggs Locality North Dakota (ND) 52 50 Wahpeton Church’s Ferry Devils Lake Rock Lake, Towner Co. Pembina Pierce Co. [Buffalo Lake] Stump Lake Eddy Co. Eddy Co. Duck Lake Lakota Arthur, Cass Co. Rock Lake Ward Co. 8 mi W Deering 2 mi NW Antler 4 mi W Deering nr Deering 12 mi W Antler 6 mi S Flaxton 10 mi E Deering 4 mi N Denbigh 5 mi E Deering 4 mi E Deering 5 mi E Deering 7 mi NE Flaxton 2 mi E Antler 3 mi NE Deering 6 mi SW Antler N of Antler 6 mi E Deering 2 mi SW Antler 5 mi SE Antler 4 mi SE Antier Antler NE Deering NE Deering NE Deering NE Deering McHenry Co. McHenry Co. McHenry Co. 3 mi SE Antler McHenry Co. McHenry Co. McHenry Co. McHenry Co. McHenry Co. McHenry Co. McHenry Co. McHenry Co. 2 mi E Loraine South Dakota (SD) 28 26 10.96 Yellowstone R. Vermilion Vermillion Vermillion 11.86 eggs/set Day/Mo 14 June 21 May 12 June 23 May 21 May 31 May 15 May 20-23 May 18 May 23 May 29 May 26 May 15 May 18 May 18 June 22 May 24 May 3 June 26 May 28 May 28 May 28 May 31 May 31 May 5 June 27 May 31 May 19 May 21 May 11 May 12 June 12 May 18 May 1 June 19 May 25 May 6 June 6 June 20 May 24 May 24 May 31 May 5 June 14 May 20 May 12 June 18 May 26 May 13 May 8 June 7? Tat 17 May 26 May 27 May # eggs 593 11 Collector JK Jensen ES Bryant WF Hill RS Judd PB Peabody F Maltby HK Job DR Ducks DR Ducks NA Francis NA Francis AD Doerge RS Judd EJ Booth GC Withey GA Withey GC Withey GC Withey D Ogg GF Abbey GC Withey GC Withey GC Withey GC Withey GC Withey GF Abbey GL Davy GA Withey GL Davy GA Withey GA Withey GL Davy GA Withey GA Withey GA Withey GC Withey GC Withey GC Withey GC Withey GC Withey GC Withey GC Withey GL Davy GC Withey GC Withey GC Withey GC Withey GC Withey GC Withey GC Withey GC Withey GC Withey FV Hayden GS Agersborg HB Bailey HB Bailey Vol. 116 Institution Continued 2002 HOUSTON: GREATER PRAIRIE-CHICKEN ON THE CANADIAN PRAIRIES 9 TABLE 1. Continued # sets sets eggs/set c eggs Locality Day/Mo Year # eggs Collector Institution Vermillion 30 May 1882 8 HB Bailey AMNH Vermillion 30 May 1882 a HB Bailey AMNH Flandreau, Moody Co. 2 June 1882 1] HB Bailey WFVZ Egan, Moody Co. 16 June 1882 ? (via BF Goss) MPM Vermillion 16 June 1882 8 HB Bailey AMNH Vermillion 23 June 1884 10 GS Agersborg CAS Roswell, Miner Co. May 1888 12 FA Patton UMA Sanborn Co. 7 May 1889 10 FA Patton WFVZ Miner Co. May 1889 11 FA Patton AMNH Roswell, Miner Co. 14 May 1889 12 FA Patton HSU Miner Co 17 May 1890 12 A Hewitt WFVZ Harrison 18 May 1890 10 WC Colt MVZ Miner Co. 10 May 1891 14 FA Patton USNM Harrison 25 May 1891 12 WC Colt MVZ Roswell 2 June 1891 11 BV Jones BMNH Bryant 20 May 1895 15 HE Lee USNM Watertown 25 May 1895 10 GW Dixon WFVZ Sanborn Co. 14 May 1898 15 FA Patton WFVZ Hamlin Co. 13 May 1903 15 HE Lee OMNH Hamlin Co. 16 May 1903 14 HE Lee MVZ Rondell, Brown Co. 18 May 1905 10 J Collins WFVZ Armour, Douglas Co. 6 May 1911 10 A Walker WFVZ Sanborn Co. 20 May 1914 — FA Patton Earlham Sanborn Co. 1 June 1922 15 FA Patton UAM Grand Total 131 119 11.10 1321 Notes: In 7th vertical column, ** indicates that a single egg was obtained from an oviduct. Egg numbers in parentheses, from incomplete sets, are not counted in totals. In 8th vertical column, ne = eggs no longer extant; otherwise, ne means northeast. In 10th vertical column, WFVZ* indicates set came from Cumberland Science Museum, Nashville Tennessee.In some clutches in some collections, some of original eggs are no longer extant, but are still counted: e.g., Stony Plain, Alberta, there are now only 11 eggs remaining from original set of 14. Institution abbreviations in 8th vertical column can be found alphabetically in Acknowledgements. H. H. Mitchell reported it as “fairly common resident in transition zone, apparently extending its range northwestward,” and Bradshaw’s annual report of the game commissioner said it was now “reported from almost all points where the [Sharptail] occurs.” Andrew Holmes reported “several fairly large cov- eys” at Davis and Henribourg, south of Prince Albert, in the fall of 1924 (Bradshaw 1925). Spread into Alberta (Figure 3) No reliable dates exist for the arrival of the Greater Prairie-Chicken in Alberta; it was never common there. Whether the “old-timers” correctly remembered it in the late 1890s, as told to Rowan, seems highly doubtful (Mitchell 1959). However, by 1911 and 1912 it was “common” in the Hanna dis- trict (Mitchell 1959), and several were shot in east- ern Alberta in the fall of 1913 (Lawton 1913). Thus in 30 years it had spread west 1000 km from Winnipeg, at an average rate of about 33 km per year, doubling the speed of its Minnesota - North Dakota spread. The first specimen from the Red Deer area was shot near Buffalo Lake, east of Mirror, by George Cook on 26 December 1914 (Horsbrugh 1918). Single individuals were shot near Bawlf in 1916 (Farley 1932), in the Delia area north- east of Drumheller in 1918 (Mitchell 1959), and south and east of Camrose in October 1926 (Farley 1932). It was hunted near Calgary and Edmonton, and seen north to Lac la Biche (Rowan 1926; Salt and Salt 1976). It nested near Buffalo Lake. Three nests were found at the south end of Beaverhill Lake, 17-18 May 1924 (Salt and Wilk 1958; Farley 1932; Mitchell 1959; Lister 1979). Three or perhaps four booming grounds near Beaverhill Lake were in use in 1925. William Rowan’s field notes show it was also common at Gough Lake and east of Big Valley, 1920-1925. In a letter to George J. Mitchell dated 15 April 1957, Rowan told of “breeding populations”’at Sullivan Lake in 1924, and also at Wavy Lake east of Daysland. Salt and Wilk (1958) mapped only nine known localities of occurrence in Alberta. Stable period in Saskatchewan, 1910-1920s The open, fertile plains, quickly broken and plant- ed to wheat and other grains, interspersed with sum- —_—— wee «ee eee ee 10 THE CANADIAN FIELD-NATURALIST TABLE 2. Main Collectors of Greater Prairie-Chicken egg sets Northern Great Plains Collector #sets State, Prov Years G. C. Withey 24 ND 1917 1934 C. P. Forge 16 MB 1900 = 1905 G. A. Withey 7 ND, SK 1927 1929 G. L. Davy 6 ND, SK 1925; 1927 F. A. Patton 7 SD 1888 1922 H. B. Bailey 6 SD 1882 G. F. Abbey 4 ND, SK 1924 1925 H. H. Pittman 3 SK 1923'' 1925 First egg set, 1879, Vermilion, South Dakota, G. S. Agersborg. Last set, 1934, McHenry Co., South Dakota, G. C. Withey, 43 sets in Western Foundation of Vertebrate Zoology. 50 museums and university collections contacted. 36 had Northern Great Plains egg sets. merfallow every second or third year, soon offered insufficient cover for Greater Prairie-Chicken in the flattest and most intensively farmed areas. Regions that remained attractive were grassy, low- lying, somewhat moist areas such as those around Old Wives Lake, the north end of Last Mountain Lake, and the Quill Lakes, together with areas in the Qu’Appelle Valley. H. H. Mitchell sighted them at Craven on 15 February 1916 and at Imperial Beach on 24 October 1922 and collected specimens at Imperial Beach on 27 October 1922 (two), and 21 October 1924. H. Eutenner (Euteneir?) shot one at Holdfast in 1920 and presented it to Mitchell at the museum; a specimen from R. A. McEwen at Nokomis had no date. (Belcher 1961). They were still present southwest of Snipe Lake until 1926, when Elizabeth [Hubbard’s] family moved to Grenfell (Hubbard 1976). Several birds were seen near Big Quill Lake at Dafoe on 10 and 11 October 1930 by Bradshaw and Bard (Belcher 1961). Four were seen at the north end of Last Mountain lake by F. G. Bard on 5 June 1936 and J. D. Soper saw a sin- gle bird at Watrous on 5 July 1936 (Soper 1970). In the Weyburn and Virden map sheet areas in southeastern Saskatchewan, they were “common and widespread” from 1910 to at least 1920. At Percival, John Nelson’s oldest brother shot one about 1928. Ralph Stueck at Abernethy and Ken Skinner at Katepwa Lake saw them regularly in the 1920s but they disappeared in the late 1930s (Callin 1980). R. D. Symons (1967) observed a booming ground near Moosomin, presumably in the 1920s. In the Moose Jaw area, sportsmen found them common through at least the early 1930s. Hugh McCrea had sightings at Old Wives Lake in the early 1930s and Tom Beveridge had them at Mortlach in 1934, where they persisted until at least early 1942 (Knight 1967). In the Allan Hills, where he farmed in the upper Arm River valley northeast of Vol. 116 Davidson, 1920-1925, R .D. Symons heard the “booming, hollow note ... in those crocus-sweet spring mornings ... awakened ... from sleep at day- break” (Symons 1967). Along the United States boundary a few were still present in 1927: one near Altawan, 8 August, several at Lonesome Butte, south of Lafleche, 9-17 September, and two at Buffalo Gap south of Willowbunch on 10 September (Soper 1970). Montana If this species indeed migrated south (or south- east?) in winter and if the 1965 and 1972 sightings in extreme southern Alberta originated from Montana, one would expect records across northern Montana, south of Alberta and western Saskatchewan. Such records do not exist. Saunders’ state list in 1921 recorded only a single specimen from a most unex- pected location, deep within the state and not even in good prairie chicken habitat (Bob Eng, personal communication), shot by John R. Bane near Huntley, northeast of Billings, in the fall of 1917 (Saunders 1921). P. D. Skaar (1987) doubted the validity of this record, believing that Saunders had not seen the specimen. If valid, might it have reached Huntley on a railroad flat car? Only in the extreme northeast corner of Montana (Sheridan County) have there been credible records at any time. The first record is for the sandhills 40 km north of Culbertson, west of the present hamlet of Medicine Lake, in 1904, when J. B. Lyons shot over 200 ‘squaretails’ in four days. A hunter shot two south of the hamlet of Homestead, 10 km farther south, about 1919. Joseph A. Morin saw them in the Wolf Creek area southwest of Plentywood in the fall of 1936. That year, William T. Krummes, assistant manager of the Medicine Lake National Wildlife Refuge (NWR), listed them as rare, but included a photo of a booming male just north of Medicine Lake. Frank Reuter, a rancher near Medicine Lake, had a large booming ground in a hay meadow until about 1940-41. Farther south, in the Yellowstone valley near the hamlet of Intake between Glendive and Savage, Art Suckow gave a description of seven or eight seen in early June 1944, the last credible record in Montana (Walcheck 1980). A careful sur- vey of Sheridan County by Bob Eng (personal com- munication) in 1952 revealed no sight or sound of Greater Prairie-Chickens, but after a hiatus of 23 years there was a single unconfirmed report in 1961-62. North Dakota In North Dakota, the arrival and distribution are well documented, after it had “followed the plow ... five hundred miles ... in fifty years ... an average of 2002 ten miles a year” (Partch 1973). The Greater Prairie- Chicken was first reported at Grand Forks in 1880, and had a “booming ground” in the most northeaster- ly county, Pembina, by 1882. By 1884, it was com- mon at Fargo and had moved 100 km west of there, a spread of 500 miles (800 km) in 50 years. By 1900, this species had spread westward almost exactly to the Montana boundary; it was 40 km short of the Montana boundary at Medora; 27 km from it at Williston; 8 km from it at Marmarth and 3 km from the boundary at Beach (Walcheck 1980); at that time it occupied all but the extreme southwestern corner of North Dakota. Between 1917 and 1934, enthusiastic oologists, David Ogg, his son-in-law, George L. Davy, and his friend George A. Withey, collected 37 clutches in McHenry and Bottineau counties, especially near Antler, North Dakota, which is just a few km south of the junction of Manitoba and Saskatchewan with the 49th parallel (Table 1, Bechard and Houston 1984). As late as 1940, roadside counts gave an esti- mated North Dakota state population of 430 000 individuals, falling to 4,000 to 5,000 by 1964 and fewer than 400 by 1972 (Stewart 1975). On the Arrowwood NWR, 360 Greater Prairie-Chickens and 2100 Sharp-tailed Grouse were counted in 1941, dropping to zero prairie chickens and 250 Sharp- tailed Grouse by 1966 (Kirsch et al. 1973). “When the proportion of grain to grassland was about 20 to 80, the Pinnated moved in and began to increase ... As the proportion moved up to 50:50, the population ... Increased to tremendous figures ... But then the ratio tipped over the other way, and there was more land in crop than in native grassland. Then the pin- nate could find plenty of food but no nesting cover ... or escape cover. He was literally plowed out of house and home” (Anonymous 1953). By 1997, the spring breeding population in North Dakota was “about 300 with most living on or near the Sheyenne National Grassland” (Kobriger 1999). Saskatchewan breeding records (including flightless young) The first Saskatchewan nest in 1897 has been mentioned above. H.C. Grose collected a set of 10 eggs near Melville on 10 June 1918 (Houston 1993). Pittman at Wauchope found nests with eggs in three successive years, 1923, 1924 and 1925 (Nero and Lein 1971). The enthusiastic oologist group residing in Antler, North Dakota, within 3 km of the Canadian boundary, collected at least two sets within Saskatchewan, and four other sets that were either within Saskatchewan or within one mile of it: 11 eggs on 6 June 1925, 9 eggs on 29 May 1927, 10 eggs on 8 June 1927, and 10 eggs on 14 June 1927 (Table 1). The sets definitively named as from with- in Saskatchewan, 12 eggs from North Portal on 3 May 1925, and 14 eggs south of Gainsborough on 12 HOUSTON: GREATER PRAIRIE-CHICKEN ON THE CANADIAN PRAIRIES 11 May 1925, were both collected by George Abbey (see Table 1 and also Bechard and Houston 1984). At Quill Lake, five chicks were seen 18 June 1929 by Fred Bradshaw, and a female with a brood was seen there by Todd (1947), 20 June 1932. J. P. Maywood found a nest with 14 eggs, 6 km south- west of Nipawin, 22 June 1947. M. G. Street flushed a female with seven young near Nipawin on 28 July 1947 (Houston and Street 1959). Wallace Black had several nest records at Somme, 1920—1942 (Hooper 1992). Peter McLellan found a hen with three young in deep grass 3 km east of Arcola on 13 September 1955 (letter, 13 September 1955). The latest report was an adult with eight half-grown young in a long grass meadow in Cypress Hills Provincial Park, 14 July 1966 (Pegg 1967). Migration? Did some Greater Prairie-Chickens migrate? Salt and Wilk (1958) explained the absence of winter sightings in Alberta (but note the 26 December 1914 specimen above) with the remarkable but unsubstan- tiated claim that Greater Prairie-Chickens “left the province during the winter.” There is reasonable evi- dence (Cooke 1885) that females, especially, migrat- ed from southern Minnesota and northern Iowa to southern Iowa and northern Missouri. However, Hamerstrom and Hamerstrom (1973) found no evi- dence of migration in their local Wisconsin popula- tion. Museum egg data sets Because of the paucity of other breeding records in the five jurisdictions, my wife Mary joined me for a week in the Western Foundation of Vertebrate Zoology, transcribing 3660 egg data sets for all Northern Great Plains species, including 34 Greater Prairie-Chicken sets. We also transcribed data in the Saskatchewan Museum of Natural History (SMNH) and the University of Saskatchewan Biology Museum (USask). Next, I wrote to all museums reported by Kiff and Hough (1985) to hold egg sets, and collated their data (Table 1). Museum egg data are the best breeding evidence extant for many localities in many years. They also provide average numbers of eggs per set: 11.5 in Alberta (n = 2 ); 10.9 in Saskatchewan (n = 8 ); 10.0 in Manitoba (n = 30); 11.9 in North Dakota (n = 50); and 11.0 in South Dakota (n = 26) (Table 1). Relative numbers Relative abundance has several times been stated in terms of the numbers of their nearest related species. In 1886, near Winnipeg, Greater Prairie- Chickens were about half as common as the Sharp- tailed Grouse (Nash, in Thompson 1890). Near Saltcoats from 1910 to 1920 they were as common as the Sharp-tailed Grouse (Baines 1956), a claim 12 THE CANADIAN FIELD-NATURALIST made also by Ferry (1910) at the Quill Lakes in 1909. In southern Saskatchewan, they were the “commonest upland game bird ... the real open prairie dwellers” at some unspecified time after 1905 (Taylor 1955). Rutherford (1914) claimed that “grouse, pinnated, ruffed and sharptail, are to be had in immense numbers.” In adjacent North Dakota, E. T. Judd (1927) found that the Greater Prairie- Chicken was “practically unknown” in the early 1890s, but increased after 1910 to become common by 1915: “if it continues to increase as it has in the past five years it can soon be classed as an abundant bird,” to a large extent replacing the Sharp-tailed Grouse. Near Raymore, Saskatchewan, it increased rapidly in three years to become equivalent in num- bers to the Sharp-tailed Grouse about 1920 (Charles Harris in W. C. Harris, unpublished manuscript). Ten-year population cycle Greater Prairie-Chicken did not persist long enough on the Canadian prairies for a population cycle to become evident. Elsewhere, Minnesota pop- ulations peaked in 1863, 1871, 1880-81, and 1894-95, and a century later in 1982 and 1992; in Wisconsin they peaked in 1940, 1950, and 1970-71 (Svedarsky et al. 1997). This approximate ten-year cycle is similar to that of the Great Horned Owl, Snowshoe Hare, Lynx, and to that of other upland game birds, the Ruffed Grouse, Sharp-tailed Grouse and Spruce Grouse elsewhere in North America (Houston et al. 2002, in press). Habitat requirements “..wildlife abundance or scarcity is fundamentally a function of habitat suitability. ... nest-brood cover [is] the weak link in the Prairie Chicken life chain” (Olson in preface to Hamerstrom and Hamerstrom 1973). Roberts (1936) postulated that the presence of grain for food was the stimulus for the westward and northwestward spread within the United States; the Greater Prairie-Chicken entered new areas soon after grain was planted. The importance of grains was cor- roborated by a 1962 study in Missouri, where 44% of diet volume was composed of corn, soybeans and sorghum, in descending order of importance, with wheat and oats together contributing another 9% (Korschgen 1962). Macoun (1900) described it as a “true prairie bird as observers speak of it always being found in the open even in the severest weather.” Everywhere it tended to choose low-lying areas with high grass. Schroeder and Robb (1993) point out that its pre- ferred habitats tend to be interspersed with or sur- rounded by oak trees, reminiscent of the Passenger Pigeon (Ectopistes migratorius). As it followed the plow westward, the Greater Prairie-Chicken presumably had the best of both worlds, but only for a short while. Long grass Vol. 116 offered cover and grain fields provided food, the two habitats often “interdigitating” (Johnsgard and Wood 1968). They seemed to take advantage of the thick grass that grew luxuriantly between the time of less- ened grazing pressure due to the demise of the Bison and the subsequent secondary over-hunting of Pronghorn (Antilocapra americana), and the later increase in cattle (graphed by Johnston and Smoliak 1976). The grassland range was “not fully stocked until about 1920.” Suppression of grass fires by set- tlers was a further benefit. Next, as even more land was cultivated, especially when cultivation reached 60% of the land area, and as ever-fewer parcels of grass interspersed with rose- hips (Judd 1905) had sufficient area to support a group of prairie-chickens, this species left (Rue and Allen 1973). And, in the absence of fire, invasion of grasslands by trees and shrubs made grassland frag- ments even smaller. In North Dakota, the ideal habi- tat was grass 20 inches (50 cm) in height and suffi- ciently dense to completely conceal a nesting female; fields annually grazed, annually hayed, or idle >10 yr were undesirable (Kirsch 1973). In Oklahoma, Jones (1963) noted that mean height of vegetation in booming grounds was 15.1 cm., at nest sites 45 cm (range 25 to 70 cm), and escape cover, 60 cm. The drought of the 1930s, complicated by over- grazing, sounded the final knell; few coveys could find luxuriant grass to provide adequate cover on the Canadian prairies. Between 1920 and 1940 there was also a slowly increasing shift from harvesting with threshing machines (producing attractive straw- stacks) to combines (Svedarsky et al. 1997); the elimination of strawstacks was not complete until 1960, after the demise of the Greater Prairie- Chicken. Decline in Alberta In Alberta, Greater Prairie-Chickens had “practi- cally disappeared” by the mid-1930s. The last sight- ing in the Drumheller area was by W.R. Salt north of Hand Hills, east of Drumheller, in 1925 (Kondla et al. 1973). They disappeared from the lek at the south end of Beaverhill Lake about 1929 (Farley 1932), except for a single in May 1932 and three seen there 24 May 1934 (Lister 1979). Decline in Saskatchewan In Saskatchewan, during the First World War all grouse numbers, including those of the Sharp-tailed Grouse and Ruffed Grouse, collapsed to such a degree that the season was closed for Sharp-tailed Grouse and Greater Prairie-Chicken from 1916 through 1920. Neil Gilmour, Game Guardian at Moose Jaw, reported in 1923 that the Sharp-tail and the Ruffed Grouse had rebounded and continued to increase, but the Pinnated, once fairly common, was 2002 not coming back in numbers as the other two species had done (Bradshaw 1923). The decline in Greater Prairie-Chicken numbers began early. They were common north of Bladworth until about 1918, with two booming hills of 12-15 birds each (Roy 1996). They were last observed booming on knolls near Langbank in 1924—1927 by Geoffrey Hewson (Belcher 1961; Hewson 1977), and last seen near Sheho about 1926 by William Niven (Houston 1949). They disappeared from the Storthoaks area by the late 1920s, certainly by 1935 (Stelfox 1980; Ray Barber, personal communica- tion). Last “booming ground” at Cut Knife was in 1931, and the last in a hunter’s bag at Maidstone was in 1932 (Symons 1967). Near Davidson and Imperial, Saskatchewan, they were “greatly decreased in numbers” by 1932, although there were still three booming grounds near Davidson and Stalwart and east of Imperial; two were seen at “Devil’s lake” west of Simpson, and five specimens were collected at Last Mountain Lake (Todd 1947). They disappeared from the Qu’ Appelle valley in the late 1930s (Callin 1980). The last observation at Raymore was of a single male observed by A. Cameron dancing with Sharp-tailed Grouse in the springs of 1947 and 1948 (Wayne C. Harris, unpub- lished manuscript). There are only four known observations within the Saskatoon area. A mounted specimen in the pos- session of N. Loucks, Main Street, Saskatoon, had been collected about 1 February 1925 (interview with J. B. Gollop about 1970). A male specimen in the University of Saskatchewan Biology Museum, was collected at or near Saskatoon, 12 October 1931 (Smith 1968). From 1935 to 1945, Judge A. H. Bence (1945) observed the “odd covey” north of Saskatoon and he once watched the courtship “booming” display of the male near a slough between Borden and Radisson. In 1946, while col- lecting specimens for the Royal Ontario Museum, Farley Mowat (1946) flushed one near Dundurn, on the road to Proctor Lake, on 21 May [date obtained from Mowat’s field notes]. We can only guess how common Greater Prairie- Chickens were in extreme southern Saskatchewan, but a few persisted until 1940 and 1941. Soper saw one or two every day at Rock Creek, south of Lafleche, on 7 and 8 June; one near Wood Mountain on 9 June 1940, and one along the Frenchman River below Val Marie on 16 June and one farther west at Middle Creek on 20 June 1941 (Soper 1970). Probably Greater Prairie-Chickens were never common near the northern edge of their maximal range, though paradoxically this was where some of the final sightings occurred; here the requisite long grass persisted in clearings along the fringe of low- lying areas in mixed forest. The last sighting at Prince Albert was about 1932 (Brooman in Houston HOUSTON: GREATER PRAIRIE-CHICKEN ON THE CANADIAN PRAIRIES 13 and Street 1959). Maurice Street or his brother Stanley saw a flock of nine, 8 km east of Nipawin, 10. -December) 1935) Mhey)weren regulars at Kelvington, Lintlaw and Somme until 1934, with the last sighting at Somme about 1942 (Hooper 1992). Several were seen along the west boundary road of Prince Albert National Park on 14 July 1940 in an area with small patches of prairie or grassy wood- land glades ... a surprise (Soper 1952). James McCunn, on his farm a mile east of Codette, saw and shot his first on 20 September 1945. Two months later, on 20 November 1945, McCunn saw a flock feeding at a granary 43 km east of Nipawin (Houston and Street 1959). At Emma Lake, a flock of 12 in natural or “moose” meadows in 1937-38 decreased to three in 1939, the year that Mowat saw single individuals in Prince Albert National Park and at Montreal Lake (Mowat 1947). The last record in central Saskatchewan was of a lone bird 5 km north of Snowden, 10 June 1950 by Walter and Billy Matthews and three other observers (Houston and Street 1959). In southern Saskatchewan, Soper’s last sightings were one near Reed Lake, just south of Morse, on 16 July 1946. a pair near Redvers on 8 August 1946; two south of Bender post office, south of Whitewood, on 10 August 1946; and one near Red Jacket on 20 September 1947 (Soper 1970). Decline in Manitoba | Having peaked in numbers, the decline was evi- dent early at Birtle, Manitoba, where it was only “abundant until 1916” (Bird 1930). Bruce Noton (personal communication) remembers seeing two in the ditch en route to Royal School, 6 km west and 2 km north of Boissevain, about 1931; his friend, Herb Patterson, shot two south of Wawanesa in the early 1930s. In the winters of 1933 and 1934, Victor Latta reported a single bird near Shelley. T. Schindler shot his last one near Clandeboye about 1934 (Gardner 1981). G. W. Malaher saw over 300 within 100 km of Winnipeg as late as 1936 (Hamerstrom 1956). Douglas Shanks told of small numbers and at least one booming-ground near Old Pinawa in the late 1930s; his last sighting was of four in the same area in the winter of 1941-42 (Taylor 1983). The Greater Prairie-Chicken disap- peared from Lyleton in the 1940s (Knapton 1979) and from the Oak Hammock Bog in 1946 (P. Romanic in Gardner 1981), but was present along the south edge of Delta Marsh “until 1950 when it declined rapidly.” (Hochbaum 1971). Three were seen at Windfield Swamp near Stonewall by Robert E. Jones in the winter of 1950-51 (personal commu- nication). John H. More saw a Pinnated Grouse with a group of Sharptails near Russell, on 8 October 1953 (letter, 19 October 1953). 14 THE CANADIAN FIELD-NATURALIST Causes of decline Bird (1961) ascribed the decline to two factors: over-shooting and destruction of nesting sites by intensified agriculture. A review article reported “It is ironic that the Greater Prairie-Chicken at first prospered from an expanding agriculture ... [until] the further development of cereal farming and cattle ranching eliminated the tall grass vegetation on which the species had depended....the late 1870s and early 1880s were wetter than normal and grass grew luxuriantly” (Johnston and Smoliak 1976). Godfrey (1986) summarized the situation similarly: “In the interim between the disappearance of the bison from the prairies (about 1880) and the beginning of inten- sive settlement (about 1920), the natural prairie grasslands flourished as never before and probably never will again. Natural grassland is ideal for the Greater Prairie-Chicken and it too flourished in that period and declined rapidly when its habitat was destroyed by the settlements of man.” Christisen (1969) has provided a list of factors detrimental to this species, with special reference to the United States: (a) conversion of grass to tilled crops, (b) removal of grass cover by grazing and haying, (c) overgrazing, (d) extensive burning, (e) loss of open grasslands to shrubs and trees, (f) drought in the 1930s, and (g) lack of winter food. Remaining populations in the 1960s were confined to areas with some Big Bluestem Grass, and in Missouri required areas with 25 to 30% grass. Another factor contributing to decline in Illinois was the decrease in production of redtop, Agrostis alba, which gave excellent nesting and brood cover, but was largely replaced by legume hays which were often harvested at the time when broods are hatching (Yeatter 1963). Depredations by raptors, an ever- present threat, seem not to have been a serious factor in the decline (Berger et al. 1963). Burning was not always detrimental; use of grassland increased the second year after a spring burn (Westemeier 1973). Robel (1970) reported from Kansas that of 60 Greater Prairie-Chicken males on three booming grounds, 2.7 km apart, only 33 took up territory and only six (two at each booming ground) in any one year were involved in mating. Whether or not such social systems are dysfunctional in declining popula- tions seems not to have been studied. Skinner (1974) found that the two small colonies persisting in northern Missouri in 1973 had suffered mainly from conversion of grassland to cropland and intensive grazing on what pastures remained. Winter and summer, prairie-chickens preferred cover of 8 to 10 inches in height, hence Skinner suggested as a management technique for this species that the remaining “warm season grasses” should “never be grazed below ten inches”! Ryan et al. (1998) found that contiguous prairie > 65 ha supported a stable population over 27 years in southwestern Missouri, Vol. 116 - whereas nest success was lower in the prairie mosaic landscape and the population there declined. Interbreeding/hybridization None of the authorities quoted above mentioned hybridization as a contributing factor to the decline. However, hybridization with the Sharp-tailed Grouse (Evans 1966; Rowan 1926; Sparling 1980; Lumsden 1970), and less commonly with the Ring-necked Pheasant (Lincoln 1950), has had deleterious effects. On Manitoulin Island, Ontario, colonized by Sharp- tailed Grouse from the north, and by Greater Prairie- Chicken from the south (Lumsden 1949), hybridiza- tion seems to have led to the complete disappearance of the Greater Prairie-Chicken (Lumsden 1970; Godfrey 1986). Grouse interbreed remarkably freely. Alberta’s first two hybrids of Greater Prairie-Chicken with Sharp-tailed Grouse were reported from Gough Lake in 1918 and from near Edmonton in 1925 (Rowan 1926). C. G. Harrold (1933) identified two hybrids with a party of Sharptails near Old Wives Lake in May 1922, but Saskatchewan’s first recorded hybrid specimen was not taken until 22 October 1933 by J.C. Lusted south of Regina at Truax (Belcher 1961). Some, perhaps most, of the last records of Greater Prairie-Chickens in the prairie provinces were in company with Sharp-tailed Grouse or Sage Grouse. At Raymore, Wayne C. Harris (personal communication) saw a hybrid Sharp-tailed Grouse x Greater Prairie-Chicken on 25 June and again on 24 December 1971. On 20 April 1987, Wayne C. Harris and Don Weidl saw a female Greater Prairie- Chicken on a Sage Grouse dancing ground between Killdeer and Val Marie in the west block of the nascent Grasslands National Park (Gollop 1987, Harris et al. 1987). Three days earlier they had seen what was hypothesized to be the first-ever hybrid Sage Grouse X Greater Prairie-Chicken. However, Dale Hjertaas (1995) photographed a Sage Grouse x Sharp-tailed Grouse hybrid south and west of Lafleche in May 1988, a hybrid similar in appear- ance to that reported earlier by Harris farther south. Similar hybrids have been reported from Montana (Eng 1973) and Alberta, with DNA analyses as proof for the latter (Aldridge et al. 2001). No Sage Grouse X Greater Prairie-Chicken hybrid specimen has yet been collected, probably due to the minimal overlap in their distribution. In Nebraska, extensive surveys in the 1960s showed that up to 17% of display grounds were mixed, and that the minimum rate of Greater Prairie- Chicken hybridization varied from 0.3 to 1.2% (Johnsgard and Wood 1968). In extreme northwest- ern Minnesota, 3 of 11 stable display grounds were mixed, and up to 3.5% of the birds were hybrids (Sparling 1980). 2002 We have no evidence incriminating the introduced Ring-necked Pheasant (Phasianus colchicus) with the decline of the Greater Prairie-Chicken in Canada. Widespread release of pheasants from the Beaver Creek Pheasant Farm south of Saskatoon, Saskatchewan, began only in 1946, after the demise of the prairie chicken. In Illinois, the relationship is more compelling: “the closure of hunting seasons on Greater Prairie-Chickens coincided with the initia- tion and expansion of pheasant stocking programs. ..remnant flocks exist outside the contiguous range of the pheasant.” Further, parasitism of prairie chick- en nests by hen pheasants was observed each year and parasitized nests were less successful than non- parasitized nests. With mixed clutches the pheasants hatched a day earlier, and the prairie chicken hen would leave “with the parasitic brood before her own eggs hatched.” A single cock pheasant would domi- nate the entire booming ground. Removal of the offending cock merely allowed a subordinate succes- sor pheasant to assume the same territory (Vance and Westemeier 1979). In-breeding of isolated remnants on the Canadian prairies may have administered the final coup de grace. In Illinois, Greater Prairie-Chicken numbers declined from several million birds in the mid-19th century (600 000 were marketed in Chicago in 1873; Westemeier and Edwards 1987), to a small remnant of 46 survivors by 1994. The dramatic crash in num- bers is depicted on four maps; extremely low fertility and egg success of these isolated survivors was attributed to isolation and inbreeding, substantiated by a quick recovery of fertility and hatching success after 271 individuals had been introduced from large populations in three other states (Westemeier et al. 1998). Last Alberta specimens Shot 1931 between Czar and Hughenden (Mitchell 1959). Shot 1938 near Youngstown (Salt and Wilk 1958). Last Alberta sightings Mid-1930s, near the Oldman River west of Fort Macleod (Mitchell 1959). 28 August 1939, two just outside Waterton Lakes National Park, not far from the Montana border (C. H. D. Clarke in Rand 1948). 1939, early 1950s, and finally in either 1961 or 1962, near Sullivan Lake (Mitchell 1959, Moyles 1987). 1940, north of Medicine Hat (Mitchell 1959; Godfrey 1966, 1986). Through 1943, but not in 1944, Dick Beddoes had them in his unpublished Christmas bird counts at Daysland, southeast of Camrose (Beddoes 1945). On 14 September 1965, B.R. Shantz (1966) saw eight birds, 13 km west of Coutts, Alberta, and only 3 km north of the Montana boundary, but HOUSTON: GREATER PRAIRIE-CHICKEN ON THE CANADIAN PRAIRIES 15 Sadler and Myres (1976) categorized this as “questionable.” 5 March 1972, a single bird near Mountain View, southwest of Cardston (Salt and Salt 1976). Last Saskatchewan specimens Shot 30 April 1931 at Imperial Beach by F.G. Bard (Belcher 1961); Shot October 1931 by Dr. J.R. Hoag at Viewfield, northwest of Estevan (Belcher 1961); Shot 24 Oct 1940 east of Simpson by F.G. Bard (Belcher 1961); Shot fall of 1942, west of Saltcoats, by Robert Cock (Houston 1949); Shot Oct 1943 at Marienthal, southwest of Estevan, by F.N. Dunk (Belcher 1961); Shot 31 Oct 1945 near Carlyle, by R.J. Fyfe (Lahrman 1957); Shot 3 Nov 1972 near Leader (Hatch 1973). Last Manitoba specimens Shot in 1957 at Delta Marsh, in the largest remnant of true tall-grass prairie, by a group of American hunters (Lawrence St. Goddard, personal commu- nication, fide Robert E. Jones). Shot in the fall of 1961 near Langruth (Salt and Salt 1976; Godfrey 1966). Last Manitoba sightings 1972, a possible sighting in the Broomhill Wildlife Management area in southwest Manitoba (Vere Hunt Scott, fide Knapton 1979). 3 October 1978, Oak Hammock Marsh, Mr. McKillop (Gardner 1981). 1983, Spruce Woods east of Brandon, possible sight record by K. Leavesly (Minish 1987b). 25 May 1986, four near Pilot Mound, observed by Doug and Burton Collins (possibly escapes from releases made at Grand Forks, North Dakota, near the same time; Robert E. Jones, personal commu- nication) Last Saskatchewan sight records Five unpublished Saskatchewan records resulted from my 1956 appeal, including the report of Rayner (and More’s from extreme western Manitoba), above. Melvin Adair (letter, 12 March 1956), east of Outram, 18 km west of Estevan, believed that Ring- necked Pheasants, Sharp-tailed Grouse and Greater Prairie-Chicken all suffered severely in the blizzard of 1948. When in the spring of 1949 he dug out his coyote traps from beneath the melting drifts that had been 7.6 m deep, he found the corpses of a covey of Pinnated Grouse that had perished in that snowdrift. His Ring-necked Pheasant count locally dropped from 169 to 1, presumably the result of the same storm, reputed to have been the worst since 1888. 16 THE CANADIAN FIELD-NATURALIST Peter McLellan of Arcola (letter, 24 March 1956) hunted Greater Prairie-Chickens in the Warmley area, 18 km north of Kisbey, until about 1936, about equal in numbers with Sharptails, and then saw a sin- gle individual there about 1946. His brother, J. S. McLellan, found them regularly in section 31-8-4w?2, 6 km north and 3 km west of Arcola, but a careful search revealed none there in 1955, the year that Peter found three young closer to Arcola. Four southern Saskatchewan sightings were near lakes. Near Lucky Lake, the last sightings of individ- ual birds were on 15 August 1943 and 16 August 1947 (Roy 1996). Five were seen at Good Spirit Lake, 17 January 1946 by J. A. Gunn (“‘jottings” given to the late Mrs. Isabel Priestly, now in posses- sion of CSH). Three birds were seen on the Otthon flats, southwest of Yorkton, early November 1946, by W. D. Lightbody (Houston 1949). J. E. Glennie found them northwest of Govan, within a mile of the east shore of Last Mountain Lake, about 1950 and 1951 (letter, 21 May 1956). Farther south, David MacDonald of Manor saw one in the spring of 1954, three in 1958 and one at a Sharp-tailed Grouse dancing ground in the spring of 1959 (Belcher 1961). Fred Sharp saw three with eight Sharp-tailed Grouse, 6 km north of Sidewood in September 1959 (Belcher 1961). Roger Tory Peterson and James Fisher, arguably the two best- known ornithologists attending the American Ornithologists’ Union meeting in Regina in 1959, saw a single cock Greater Prairie-Chicken within a mile of the east edge of Old Wives Lake on 25 August 1959 (Peterson 1960). Two were observed carefully near a highway near Stoughton in the fall of 1960 (Pringle 1961). Five were carefully described near the east edge of Old Wives Lake on 3 September 1961 (Pratt 1967). One was seen by J. Luthi, 5 km north and 13 km west of Simpson on 12 May 1962 (Luthi 1963). Twelve appeared on the farm of Gilbert Johnson near Marchwell on 16 February 1967 (Johnson 1967). A single individual was with six Sharptails near Ruthilda, mid-July 1971 (Wapple 1977). A hybrid was seen by Wayne C. Harris near Raymore, 25 June and 24 December 1971 (Houston 1971, 1972). Single birds were seen near Mortlach on 19 December 1971 and near Caron on 16 April 1972 (Brazier 1972a, 1972b). David W. Robinson saw two with Sharptails near Avonlea, 20 September 1975 (Serr 1976). Derek Kreuger saw two in the ungrazed section of the northeast quarter of Saskatchewan Landing Provincial Park in 1977 and again in 1978 (Blood and Anweiler 1979). Hjertaas et al. (1993) collected additional obser- vations for the RENEW report as follows: R. K. Brace, a biologist with Canadian Wildlife Service, saw one southwest of Lafleche about the summer of 1966. B. Campbell saw one on a Sharp-tailed Grouse lek 18 km south and 13 km east of Mortlach in April Vol. 116 - 1968. Don Jackson saw one 12 km north and 4 km east of Plunkett in October-November 1972. J. B. O’Neil saw one on a roadside 5 km east of Viscount in February 1973 and then saw a small flock on three consecutive Fridays 7 km east of Plunkett in February 1975. Gerhard Stuewe saw a group of between six and eight at Teo Lake, west of Kindersley, in mid-September 1975. Ross Campbell, a hunter, saw one and then three others within 8 km of the South Saskatchewan River, south of Riverhurst in Game Management Zone 26C in the fall of 1983, and then two near Floral in the winter of 1984-85 through 22 Feb 1985 (Hjertaas et al. 1993). Wayne C. Harris saw a female Greater Prairie- Chicken in the west block of the National Grassland Park (34-2-12w3) on 20 April 1987 (already men- tioned under “Interbreeding/hybridization, above). Then on 8 May 1987, Harris saw three Greater Prairie-Chicken males on the United States boundary just west of the west margin of the east block of the park (5-1-7w3); they flushed from an upland area and flew into Valley County, Montana, a region lacking either historic or recent records of Greater Prairie-Chicken. This was the third last report from Saskatchewan. The second last sighting of probable Greater Prairie-Chickens in Saskatchewan were convincingly described north of Big Muddy by Carol Bjorklund, 28 July 1991 (Koes and Taylor 1991). Near the soli- tary male were three Sharp-tailed Grouse hens with broods (Carol Bjorklund, personal communication). Doris Silcox saw a single individual, together with 25 Sharptails and up to 9 Gray Partridge (Perdix perdix) at Carlyle, mid December 1991 through 18 Feb 1992 (Koes and Taylor 1992); this was the final sighting recorded for Saskatchewan and the last for Canada. One cannot but wonder how the individuals sighted above escaped detection in so many locali- ties for so long without being seen, unless they fly long distances from the nearest stable groupings in the Dakotas. Should that be the explanation, then more sightings may occur in future, though without appreciable hope of re-establishment anywhere in the Canadian west. Indeed, the Greater Prairie- Chicken recovery team was disbanded in 1994. The species had been listed in Canada as Endangered in 1978, and then as officially Extirpated in 1990 (Hjertaas 1998). Maps of range The original zone of overlap between the Greater Prairie-Chicken and the Sharp-tailed Grouse was ini- tially narrow (Figure 3 in Johnsgard and Wood 1968), but as the Greater Prairie-Chicken spread westward this zone of overlap greatly enlarged. By January 1955, Aldrich and Duvall (1955) showed habitat still occupied by Greater Prairie-Chicken in southern Manitoba, but none anywhere in 2002 HOUSTON: GREATER PRAIRIE-CHICKEN ON THE CANADIAN PRAIRIES 17 Saskatchewan, and only a small remnant area left in central Alberta. By 1979, Westemeier’s map showed substantial populations persisting only in South Dakota, Nebraska, Kansas and Oklahoma. Acknowledgments I wish to thank Harry Lumsden for unpublished information on this species in northwestern Ontario, and Ken J. Lungle for copying the 27-page publica- tion (1959) by George J. Mitchell, then Game Biologist for the Alberta Department of Lands and Forests. Wayne Pepper reviewed the annual reports of the Saskatchewan Game Commissioner, 1905-1930. Carol Bjorklund offered data from her 1991 field notebook. Dale Hjertaas contacted Ray Barber of Storthoaks for more detailed information beyond that mentioned in Stelfox’s 1980 article. James R. Jowsey provided information about the Saskatchewan egg collection of Ralph and Norman Jowsey, now in the Provincial Museum of Alberta, Edmonton (PMA). Jocelyn Hudon of PMA provided information concerning the John Kowal egg collec- tion in their museum. Robert E. Jones, retired upland game specialist, provided detailed information con- cerning Manitoba sightings. Robert W. Nero provid- ed the information compiled from early newspapers in southern Manitoba, 1884-1949, prepared by Anita Steeg Kovacs. Larry Igl, Dale Hjertaas, Robert W. Nero, Allan R. Smith, and Wayne Pepper offered constructive criticism. Dan Svedarsky provided three recent publications dealing with Greater Prairie- Chicken in the northern states. Lloyd W. Kiff pro- vided unlimited assistance during our visit to the Western Foundation of Vertebrate Zoology (WF VZ) and Frieda Kinoshita kindly supplied data on sets added to the WFVZ collection subsequently. Marc Bechard recorded data sets in the National Museum of Canada. The entire project of checking egg collec- tions would not have been possible without the Inventory of Bird Egg Collections of North America, 1985, compiled by Lloyd F. Kiff and Daniel J. Hough. The following 34 curators responded to mail and e-mail requests, supplying data re egg sets: Tim Armstrong, Adams State College, Alamosa, CO (ASC); Emanuel Levine, American Museum of Natural History, New York (AMNH); Ann Kessen, Bell Museum of Natural History, Minneapolis (BMNH); Karen Cebra, California Academy of Sciences, San Francisco (CAS); Will Post, Charleston Museum, Charleston, S.C. (CM); Starla Miller, Clemson University (CU); Tim Matson, Cleveland Museum of Natural History, Cleveland, OH (CMNH); Gene K. Hess, Delaware Museum of Natural History (DMNH); Kirstie M. Bay, Denver Museum of Natural History (DM); John Iverson, Joseph Moore Museum of Natural History, Earlham College, Richmond, Indiana (Earlham); Tom Webber, Florida Museum of Natural History, Gainesville (FMG); David Willard, Field Museum of Natural History, Chicago (FMNH); Tamar Danufsky, Wildlife Department Museum, Humboldt State University, Arcata, California (HSU); Nathan Kraucanus, Milwaukee Public Museum (MPM); Allen J. Kihm and Rand Rodewald, Minot State University (Minot); Alison Pirie and Raymond A. Paynter, Jr., Museum of Comparative Zoology, Harvard University (MCZ); Carla Cicero, Museum of Vertebrate Zoology, University of California Berkeley (MVZ); Michel Gosselin, National Museum of Canada, Ottawa (NMC) [now Canadian Museum of Nature]; Joseph Bopp, New York State Museum, Albany (NYSM); Gary D. Schnell, Oklahoma Museum of Natural History, Norman, OK (OMNH); John M. Condit, Ohio State University Museum of Biological Diversity, Columbus (OSUM); Fred C. Sibley, Peabody Museum, Yale University, New Haven (PMYU); Jocelyn Hudon and W. Bruce McGillivray, Provincial Museum of Alberta, Edmonton (PMA); Selma Glasscock, Rob and Bessie Wilder Wildlife Foundation, Sinton, Texas (RBWWFEP); Ross James, Royal Ontario Museum, Toronto (ROM); the late Bob Kreba, Royal Saskatchewan Museum, Regina (RSM); Robert McKernan, San Bernardino County Museum (SBCM); Krista A. Fahy, Santa Barbara Museum of Natural History (SBMNH); Philip Unit, San Diego Natural History Museum (SDNHM); Nancy Glover McCartney, University of Arkansas Museum (UAM); Katherine L. Doyle, University of Massachusetts, Amherst (UMA); Janet Hinshaw, University of Michigan Museum of Zoology, Ann Arbor (UMMZ); Alan Gubanich, University of Nevada, Reno (UNR); William J. Maher, University of Saskatchewan (USask); Thomas C. Erdman, University of Wisconsin, Green Bay (UWGB); Frieda Kinoshita, Western Foundation of Vertebrate Zoology (WFVZ), Los Angeles, since moved to Camarillo. The following 16 institutions had egg sets of this species, but none from the Northern Great Plains: Clayton White, Brigham Young University, Provo, Utah; Arthur R. Clark, Buffalo Museum of Science, Buffalo, New York; Mary Hennen, Chicago Academy of Sciences, Chicago, Illinois; Jane E. Deisler-Seno, Corpus Christi Museum of Science and History, Corpus Christi, Texas; Christine Adkins, Cowan Vertebrate Museum, University of British Columbia, Vancouver; Steven D. Bailey, Illinois Natural History Survey, Urbana, Illinois; Kerri Leedy, Oakes Museum of Natural History, Messiah College, Grantham, Pennsylvania; Mark B. Robbins, Natural History Museum, University of Kansas, Lawrence, Kansas; Thomas E. Labedz, University of Nebraska, Lincoln, Nebraska; Albert G. Mehring, State Museum of Pennsylvania, Harrisburg, Pennsylvania; Janice Hall, Putnam Museum, Davenport, lowa; David Denton and 18 THE CANADIAN FIELD-NATURALIST Roseanna Humphrey, University of Colorado Museum, Boulder, Colorado; Gary Shugart, University of Puget Sound, Tacoma, Washington; Edward Marks, Museum of Natural History, Univer- sity of Wisconsin Stevens Point, Wisconsin. 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United States Fish and Wildlife Service, Washington, D.C. Yeatter, R. E. 1963. Population responses of prairie chickens to land-use changes in Illinois. Journal of Wildlife Management 27: 739-757. Received 18 May 2000 Accepted 10 April 2002 Status of Common Eiders, Somateria mollissima, Nesting in the Digges Sound Region, Nunavut J. MARK HIPFNER!, H. GRANT GILCHRIST?, ANTHONY J. GASTON? and DAvID K. CAIRNS* 'Biopsychology Programme, Memorial University of Newfoundland, St John’s, Newfoundland A1B 3X9 Canada Canadian Wildlife Service, Suite 301, 5204 - 50th Avenue, Yellowknife, Northwest Territories X1A 1E2 Canada. E-mail: grant.gilchrist@ec.gc.ca [Author for correspondence] 3Canadian Wildlife Service, 100 Gamelin Boulevard, Hull, Quebec K1A 0H3 Canada 4Department of Fisheries and Oceans, Box 1236, Charlottetown, Prince Edward Island C1A 7M8 Canada Hipfner, J. Mark, H. Grant Gilchrist, Anthony J. Gaston, and David K. Cairns. 2002. Status of Common Eiders, Somateria molissima, nesting in the Digges Sound region, Nunavut. Canadian Field-Naturalist 116(1): 22-25. There is little information available on status of Common Eider, Somateria mollissima, populations nesting in Arctic Canada. We surveyed Common Eiders in the Digges Sound region, Nunavut, on 19-20 July 1999, for comparison with similar surveys in 1980-1983. On six islands for which data were available in both periods, there were 4—5 times as many nests in 1999 as in the early 1980s. The mean clutch size late in incubation in 1999 (3.3 eggs per nest) was similar to that in 1983 (3.2 eggs), but lower than in 1982 (3.9 eggs); timing of hatching was similar in the two periods. All three females caught during incubation were of the subspecies S. m. borealis. In contrast to Common Eiders S. m. sedentaria nesting in the Belcher Islands, Common Eiders nesting in the Digges Sound region appear to be faring well. Key Words: Common Eider Somateria mollissima, clutch size, Digges Sound, egg size, populations, timing of hatching. Common Eiders, Somateria mollissima, are an Ivujivik, Québec, which lies in the heart of the important subsistence and cultural resource for Inuit region (Gaston et al. 1985). In this paper, we report people in Canada’s eastern Arctic, in additon to being —_ the results of our 1999 surveys and make compar- important components of arctic marine ecosystems isons with observations from 1980—1983, to assess (Reed 1986). Despite this, there is little information the status of the Common Eider population nesting available on status and trends of arctic-nesting in this region. Common Eider populations. Historical data exist for only a few locations in eastern Arctic Canada: the Study Area Belcher Islands and nearby Québec coast The Digges Sound region, and its seabird commu- (Nakashima and Murray 1988), Ungava Bay nity, was described in detail by Gaston et al. (1985; (Chapdelaine et al. 1986), the West Foxe Islands _ see Figure 1). It is dominated by a very large aggre- (Cooch 1977; Cooch 1986), the Hell’s Gate- gation (300 000 pairs) of Thick-billed Murres, Uria Cardigan Strait region (Prach et al. 1986), and the Jlomvia, that breed on the tall cliffs of East Digges Digges Sound region (Gaston et al. 1985). There is Island and the Québec mainland south and west of evidence that Common Eider, S. m. sedentaria, pop- Cape Wolstenholme. There are also many small, ulations nesting in the Belcher Islands have declined low-lying islands that support nesting colonies of sharply (Robertson and Gilchrist 1998). There also is Common Eiders and other marine birds. Potential evidence of long-term declines in S. m. v-nigra pop- _ predators on Common Eider eggs and ducklings that ulations of northern Alaska and the western we saw on these small islands included Herring Canadian arctic (Suydam et al. 2000). Updated infor- Gulls, Larus argentatus, Glaucous Gulls, L. hyper- mation on other populations is urgently required in boreus, and Great Black-backed Gulls, L. marinus. order to assess their status and devise effective con- The former two species breed in colonies throughout servation strategies (compare Reed and Erskine Digges Sound (note that there were >31 adult 1986; Dickson 1997). Glaucous Gulls with > 6 ca. 15 to 20 day old young We counted Common Eider nests on six islands on South Skerry #3 on 19 July in 1999; this colony in the Digges Sound region, Nunavut, on 19-20 __was not reported in Gaston et al. 1985). Only one July 1999. Lying at the western end of Hudson Great Black-backed Gull, a first-summer bird, was Strait, Digges Sound appears to be near the south- seen in the Digges Sound region during 1980-1983, ern limit of the breeding range of the migratory S. but there were three adult-plumaged birds near South m. borealis subspecies, and near the northern limit Skerry #1 on 19 July in 1999; we saw no sign of of the breeding range of the non-migratory S$. m. breeding. There was no evidence of predation by sedentaria subspecies (Abraham and Finney 1986). © Arctic Foxes, Alopex lagopus, on any islands that we Common Eiders are an important source of meat, visited. A single Polar Bear, Ursus maritimus, was eggs, and down for people from the community of _ present in Digges Sound in mid-July in 1999, but we pe 2002 HIPFNER, GILCHRIST, GASTON, AND CAIRNS: COMMON EIDERS IN DIGGES SOUND 23 i “304) 02 NCRTH SKERRIES Po “On 2 SOUTH sa SKERRIES GINGI ISLAND _) Ivugivik —— ISLAND NUVUK HARBOUR Scale of kilometres Figure 1. Map of the southern Digges Sound region showing the islands that were surveyed for Common Eiders in 1999. saw nothing to suggest that it had been on any islands that we visited. Methods We surveyed six islands in the Nuvuk Islands region on 19 July 1999, including some of those on which the highest numbers of nests were found dur- ing surveys in the 1980s (in particular, the South Skerries; see Table 1, Figure 1). Results of our sur- veys should be comparable to those in the 1980s, because similar techniques were used; the one exception was that the number of nests was only estimated, not counted, on South Skerry #1 in 1981 (see Table 1). In 1999, each island was surveyed on foot by a crew of 3 who walked 5 m apart. As the islands were too large to be covered in one sweep, the crew member at the outside of the line marked the extent of each pass with rocks. The crew then turned around and returned, counting nests on the opposite side of the line. In this manner, islands were completely surveyed with little risk of nests being missed or counted more than once. A nest was counted if it had a well-defined cup with down; in this region, down is placed in the nest bowl after lay- ing has been initiated (H. G. Gilchrist unpublished), so that downy nest cups without eggs represent nest- ing attempts that have failed. When a nest was found, its contents (e.g., empty, two eggs, three eggs and one duckling, etc.) were called out to one crew member who recorded the information. We thus obtained information on clutch sizes, and a crude indication of the timing of hatching. We also mea- sured the length and maximum breadth of 100 eggs in 36 clutches. We attempted to survey island groups to the north on 20 July, but rough seas and thick fog hampered our efforts. We returned to South Skerry #1 late in the afternoon on 20 July, in order to catch adult eiders. Bill measurements and blood samples were TABLE 1. Common Eider nest counts in the Digges Sound region in different years. Year Island 1981 1983 1999! South Skerry #1 fn 42 326 (255) South Skerry #2 - 19 66 (48) Pitsulak City 3 - 2 (2) Green 4 20 25 (24) Yellow ] _ 2 (2) Black 0 - 0 'Counts in 1999 are: total nests with well-defined cups (total nests with > 1 egg/duckling); no such distinction was made on counts in 1981 or 1983. > : - “Estimate only, no systematic count was made. 24 THE CANADIAN FIELD-NATURALIST taken to determine the subspecific status of these birds (see Mendall 1986). Results We caught three incubating female Common Eiders on South Skerry #1 on 20 July. Bill measure- ments indicated that all three were of the subspecies S. m. borealis (cf. Table 1 in Mendall 1986): frontal extensions ranged from 13.2 to 16.3 mm, nostril extensions 25.7 to 31.0 mm, and total bill lengths 63.2 to 66.6 mm. On the six islands we surveyed in 1999, we count- ed 421 nests with well-defined cups, of which 331 had at least one egg/duckling (Table 1). Our total count in 1999 was about 4-5 times higher than counts on the same six islands in the early 1980s (Table 1). Note that the count of 158 nests (range: 143-181) reported for the South Skerries in Gaston et al. (1985) was for all four of these islands, with the highest count from South Skerry #4 (73 nests). We did not do counts on South Skerry #3 or #4 in 1999. In nests with at least one egg/duckling on South Skerry #1, South Skerry #2, and Green Island (n = 327; Table 1), clutch sizes late in incubation ranged up to 6 eggs, with a mean of 3.33 + 1.03 (SD) eggs per nest. Common Eider clutch sizes varied among years in the Digges Sound region (ANOVA, Fy 44,= 7-53, P< 0.001). Tukey’s tests (at the a=0.05 significance level) indicated that the mean clutch size in 1999 was similar to that in 59 nests on the same three islands late in incubation in 1983 (3.20 + 1.13), but smaller than that in 59 nests late in incubation on Eider Island and the North Skerries in 1982 (3.86 + 1.09). The mean clutch size in 1999 was similar to that reported for S. m. borealis popu- lations elsewhere in Canada (means of 2.72—3.52 eggs; cf. Table 3 in Prach et al. 1986). At least one duckling was present in 0.9% of active nests (n = 331) found on 19 July, all of them on South Skerry #1 and #2. We also saw five broods on the sea off South Skerry #1 on this date, with 2-5 ducklings per brood. These observations suggest that hatching began about 17 July 1999, similar to the timing in the early 1980s: Common Eider ducklings were first seen on 17 July 1980, 21 July 1981, 14 July 1982, and 20 July 1983. For 100 eggs from 36 entire clutches measured on South Skerry #1, Green Island, and Pitsulak City, the mean length was 75.4+2.7 (SD) mm (range: 68.5—82.8 mm), and the mean breadth 49.3 + 1.3 (SD) mm (range: 46.8-53.3 mm). Eggs were not measured on surveys in the early 1980s, but Common Eider eggs in the Digges Sound region in 1999 were similar in size to those laid by S. m. bore- alis elsewhere in Canada (mean lengths: 74.0 + 3.3 to 75.8 + 3.3 mm; mean breadths: 49.6 + 1.7 to 49.7 + 1.4 mm; cf. Table 2 in Prach et al. 1986). Vol. 116 Discussion The total number of Common Eider nests that we counted on six islands in the Digges Sound region in 1999 was about 4—5 times the number counted on the same six islands in the early 1980s. It can be dif- ficult to interpret the results of surveys such as these (Robertson and Gilchrist 1998). Many Common Eiders may forego breeding in years when environ- mental conditions are unfavourable (Coulson 1984), and as a result, low nest counts in such years could be wrongly interpreted as being indicative of popula- tion declines. Similarly, surveys that detect more nests, as did ours, could be wrongly interpreted as indicating population increases if the original sur- veys were carried out in unfavourable years with extensive non-breeding. While the latter is possible, we do not believe that this was the case here. For Common Eiders, breeding parameters such as egg size, clutch size, and the timing of laying are sensitive to variation in environmental conditions during the pre-laying period (Robertson 1995). Egg and clutch sizes that we measured were very similar to those reported for Common Eiders elsewhere in the eastern Canadian arctic, and clutch sizes were similar to, or smaller than, those in the Digges Sound region in the early 1980s. Moreover, the timing of hatching and dates of observations of first broods appeared to be very similar during the two survey periods. Collectively, these results suggest that it is unlikely that conditions were so poor that only low proportions of eiders bred in all years when surveys were carried out in the early 1980s. Given that sur- vey techniques were similar in the two time periods, involving counts of all active and failed nests, we conclude that the substantial increase in the number of Common Eider nests in the Digges Sound region probably reflects an actual increase in the size of the local breeding population. If so, this raises two important questions. First, what factors might have caused the increases in the numbers of Common Eiders nesting in this region since the early 1980s? And second, if real, are increases in this region indicative of increases in Common Eider S. m. borealis populations elsewhere in eastern Arctic Canada? We do not have definitive answers to either ques- tion. Given that there were no obvious differences in environmental conditions between the two survey periods, the dramatic increases that we detected may have been due to lower levels of local human exploitation than occurred previously. Common Eider populations in the Foxe Islands declined dra- matically between 1956 and 1976, apparently due to local harvesting practices. Many eider colonies in that region are easily reached by hunters traveling by boat from Cape Dorset, and the subsistence harvest of eggs, and especially the shooting of females, had dramatic effects on the population (Cooch 1986). It 2002 may be noteworthy that the largest increases in the number of Common Eider nests in the Digges Sound region occurred on the South Skerries. These islands are off the regular travel routes, and rarely are visited by people from Ivujivik (Gaston et al. 1985). As a result, they may function as safe nesting refuges dur- ing times when few people are venturing out specifi- cally to harvest eiders or their eggs. Unfortunately, no data exist to allow us to directly examine the pos- sibility that the apparent increase was due to lower local harvests of Common Eiders, or eider products, in recent years. We also have no information with which to assess the role that immigration from local or more distant colonies might have played. Might the apparent increases that we detected in Common Eider populations in the Digges Sound region reflect increases elsewhere? Although popula- tions of S. m. sedentaria appear to have declined considerably in the Belcher Islands, an island group to the south of Digges Sound, this was probably due to severe ice conditions in Hudson Bay that caused high overwinter adult mortality in this non-migratory subspecies during the early 1990s (Robertson and Gilchrist 1998). Causes of declines in populations of S. m. v-nigra in northern Alaska and the western Canadian arctic are unknown (Suydam et al. 2000). It is both costly and logistically difficult to con- duct surveys of Common Eiders at arctic colonies, and they cannot normally be carried out on a large geographic scale. Instead, small clusters of nesting islands are usually surveyed and the results cautious- ly extrapolated to the larger population (Cooch 1986). However, surveys conducted in regions where eiders have historically been, or are currently being, subjected to local subsistence harvests cannot reli- ably be extrapolated to non-harvested populations, because local anthropogenic effects could easily swamp large-scale environmental effects. Conse- quently, repeat surveys of colonies elsewhere are now required to assess trends in S. m. borealis popu- lations in other areas. Given these considerations, it is apparent that an ideal monitoring scheme for Common Eiders would include both harvested and non-harvested populations. Although this would add substantially to the cost of monitoring programs, it should allow investigators to distinguish between the local effects of human harvesting and environmental effects that operate over a broader geographic scale. This would aid greatly in management. Acknowledgments We thank David and John Geale, as well as the residents of Ivujivik, Québec, for assistance with the surveys. The surveys were supported by the Canadian Wildlife Service, Memorial University of Newfoundland, the Natural Sciences and Engineer- ing Research Council of Canada, the Northern Scientific Training Program, and the Polar Conti- HIPFNER, GILCHRIST, GASTON, AND CAIRNS: COMMON EIDERS IN DIGGES SOUND iy) nental Shelf Project of Natural Resources Canada. We thank Scott Gilliland for insightful comments on a previous draft of this manuscript. Literature Cited Abraham, K. F., and G. H. Finney. 1986. Eiders of the eastern Canadian Arctic. Pages 55—73 in Eider Ducks in Canada. Edited by A. Reed. Canadian Wildlife Service Report Series Number 47, Ottawa. Chapdelaine, G., A. Bourget, W. B. Kemp, D. J. Nakashima, and D. J. Murray. 1986. Population d’Eider 4 duvet prés des c6tes du Québec septentrional. Canadian Wildlife Service Report Series 47: 39-50. Cooch, F. G. 1977. Changes in the avifauna of the West Foxe Islands, Northwest Territories, 1956-1976. Cana- dian Field-Naturalist 91: 314-317. Cooch, F. G. 1986. The numbers of nesting Northern Eiders on the West Foxe Islands, NWT, in 1956 and 1976. Canadian Wildlife Service Report Series 47: 114-118. Coulson, J. C. 1984. The population dynamics of the Eider Duck Somateria mollissima and evidence of exten- sive non-breeding by adult ducks. Ibis 126: 525-543. Dickson, D. L. 1997. King and Common eiders of the western Canadian Arctic. Canadian Wildlife Service Occasional Paper Number 94, Ottawa. Gaston, A. J., D. K. Cairns, R. D. Elliot, and D. G. Noble. 1985. A natural history of Digges Sound. Canadian Wildlife Service Report Series Number 46, Ottawa. Mendall, H. L. 1986. Identification of eastern races of the Common Eider. Canadian Wildlife Service Report Series 47: 82-88. Nakashima, D. J., and D. J. Murray. 1988. The Com- mon Eider (Somateria mollissima sedentaria) of eastern Hudson Bay: a survey of nest colonies and Inuit ecologi- cal knowledge. Environmental Studies Revolving Funds Report Number 102, Ottawa. Prach, R. W., A. R. Smith, and A. Dzubin. 1986. Nesting of the Common Eider near Hell’s Gate-Cardigan Strait polynya, 1980-1981. Canadian Wildlife Service Report Series 47: 127-137. Reed, A. 1986. Eiderdown harvesting and other uses of Common Eiders in spring and summer. Canadian Wild- life Service Report Series 47: 138-146. Reed, A., and A. J. Erskine. 1986. Populations of the Common Eider in eastern North America: their size and status. Canadian Wildlife Service Report Series 47: 156-162. Robertson, G. J. 1995. Annual varation in common eider egg size: effects of temperature, clutch size, laying date, and laying sequence. Canadian Journal of Zoology 73: 1579-1587. Robertson, G. J., and H. G. Gilchrist. 1998. Evidence of population declines among Common Eiders breeding in the Belcher Islands, Northwest Territories. Arctic 51: 375-381. Suydam, R.S., D. L. Dickson, J. B. Fadely, and L. T. Quackenbush. 2000. Population declines of King and Common Eiders of the Beaufort Sea. Condor 102: 219-222. Received 26 April 2000 Accepted 28 March 2002 Flathead Chubs, Platygobio gracilis, in the Upper Missouri River: The Biology of a Species at Risk in an Endangered Habitat SHANNON J. FISHER!, DAviD W. WILLIS?, MICHAEL M. OLSON?, and STEVEN C. KRENTZ3 ‘Minnesota Department of Natural Resources, Windom, Minnesota 56101 USA *Corresponding author. Department of Wildlife and Fisheries Sciences, South Dakota State University, Brookings, South Dakota 57007 USA.; e-mail: david_willis @sdstate.edu 3U.S. Fish and Wildlife Service, Bismarck, North Dakota 58501 USA Fisher, Shannon J., David W. Willis, Michael M. Olson, and Steven C. Krentz. 2002. Flathead Chubs, Platygobio gracilis, in the upper Missouri River: The biology of a species at risk in an endangered habitat. Canadian Field-Naturalist 116(1): 26-41. Flathead Chub, Platygobio gracilis, populations have declined in portions of their range where natural flood-pulses and functioning backwaters have been eliminated. Observations of these declines have led ecologists to consider both the species and their at-risk habitats. Backwater and sandbar habitats in the Missouri River, North Dakota were sampled during a high flow (1997) and an average flow (1999) year. Of the 817 larval fishes captured in the backwaters, none were Flathead Chubs. Significantly fewer (P<0.01) Flathead Chub adults and juveniles were captured in the backwaters than from sandbar habitats during all sample periods. Male chubs were all mature at 110-mm total length (TL) and age 2, whereas females were not all mature until age 3 and 170-mm TL. In 1997, Ostracoda, Hemiptera, and Copepoda dominated the diets, whereas in 1999, Coleoptera, Trichoptera, and Hymenoptera were utilized in significantly greater amounts (P<0.05). Flathead Chub maximum age was 5 and the mean back-calculated total lengths at age were 104-, 153-, 186-, and 223-mm TL for ages 1-4. Incremental growth analyses indicated that greater increases in total length were attained in 1997 than in 1999; however, body condition was significantly higher in 1999 (P<0.006). The contradiction between growth and condition cannot be explained; however, it could be energetically advantageous to maintain a more fusiform body shape during high flows. Although the backwaters were not important physical habitat for Flathead Chubs, backwater productivi- ty may have critically contributed to the prey base during the high flow period of 1997. Key Words: Flathead Chub, Platygobio gracilis, Cyprinidae, habitat alterations, food habits, Coleoptera, Copepoda, back- waters, Missouri River, North Dakota. Cross et al. (1986) and Pfleiger and Grace (1987) food resource for native fishes obligated to flowing reported that Flathead Chub, Platygobio gracilis water habitats. Eckblad et al. (1984) observed sub- Richardson, populations have decreased by as much _ stantial increases in channel invertebrate densities as 98% over portions of their historical range. Grady below connection points with floodplain wetlands and Milligan (1998) and Gelwicks et al. (1996) both and Kennedy (1979) suggested that restricted back- observed Flathead Chub declines in the channelized __ water discharge may create invertebrate voids in the and impounded segments of the Missouri River. main channel. Therefore, understanding the relation- These declines are not well understood; however, _ ships that Flathead Chub populations maintain with degraded habitats are a suspected cause (Lee et al. _ their habitats and food resources under varying water 1980). Researchers have correlated sensitive species conditions may be critical. declines with channel and flow modifications (Hesse The relatively unknown status and function of et al. 1989). In addition to the problems of popula- Flathead Chub populations in North American river tion loss and habitat modifications, Flathead Chub systems remain a point of interest. Furthermore, are difficult to study because their life history has opportunities to study Flathead Chubs and other not been thoroughly documented. native fishes in relatively unaltered habitats and Fisher (1999) noted that a segment of the Missouri hydrographs have become increasingly rare. There- River in North Dakota contained Flathead Chubs and fore, the objectives of this study were to (1) assess was maintaining historical habitat characteristics, general Flathead Chub biology, including age and including natural flood-pulse patterns from growth, food habits, habitat use, size structure, and Yellowstone River inflow and a functioning lateral maturation patterns, (2) contrast life history patterns relationship with floodplain backwaters. Scott and between two years with substantially different Nielsen (1989) and Sabo and Kelso (1991) demon- hydrographs, and (3) assess and discuss the potential strated that backwater habitats are important rearing importance of backwater habitats to Flathead Chubs. and nursery areas for numerous native fishes.. This reference information from a stable Flathead Additionally, Amoros (1991) suggested that flood- Chub population in a healthy river stretch should plain flushing transports invertebrates from backwa- help biologists evaluate populations in degraded lotic ters to the main channel, providing an important systems. 26 2002 Study Area The Missouri River study area lies between the Yellowstone River confluence and Lake Sakakawea in North Dakota (48°N, 103.9°W; river kilometers 2510-2518: river mile markers 1569-1574). This Missouri River stretch was selected because the unregulated Yellowstone River discharge provides a nearly historic flood-pulse hydrograph. Also within this stretch, several functioning backwater wetlands and numerous clusters of sandbar habitats are pre- sent. We monitored two backwaters and two sandbar complexes for use by Flathead Chubs. The two back- waters were described as off-channel habitats that connected to the Missouri River during one or more periods annually and maintained minimal flow (Fisher 1999). Surface areas for both backwaters were variable depending upon the prevailing hydro- logic conditions; however, each was characterized by annual inundation of terrestrial vegetation, silt and mud dominated substrate, presence of marsh smartweed, Polygonum coccineum, and depths that generally did not exceed 1.5 m. The sandbar com- plexes were each located in close proximity to the outlet channels of the above-mentioned backwaters and also varied in extent and persistence due to hydrologic conditions. Although the location and extent of sandbars was dynamic, the habitat was dominated by sand substrates, tended to be free from woody or other vegetative debris, and ranged in depth from 0.2 to 2.0 m. Methods Sampling Schedule and Habitat Analyses To provide a distinct comparison, we sampled Flathead Chubs in the Missouri River during one hydrograph cycle that approximated mean flow (1999) and during another cycle that included sub- stantially greater flow volume (1997; USGS 1999*; Figure 1). In 1997 and 1999, temperature, turbidity, and dissolved oxygen data were collected from chan- nel-sandbar and backwater habitats. See Table 1 for sampling schedule information. We were unable to sample sandbars during sample period 2 in both 1997 and 1999 due to high waters from mountain FISHER, WILLIS, OLSON, AND KRENTZ: FLATHEAD CHUBS IN UPPER MISSOURI 27 Day of year FIGURE 1. Cumulative daily flow (m3/sec) for the Missouri River below the Yellowstone River confluence and above the Lake Sakakawea headwaters in North Dakota for calender days 75-260 of 1997 and 1999. The solid line represents the total sum of the flow estimates for gauging stations in the Missouri River and Yellowstone River upstream from the conflu- ence. The shaded area represents the 40-year mean flow for the period 1959-1999. snowmelt that inundated the habitat from mid-May through mid-June. Data were tested for normality with a Shapiro-Wilkes test (UNIVARIATE proce- dure; SAS 1990). Likely due to small sample size, not variance, none of these data sets were normally distributed; therefore, a nonparametric Kruskal- Wallis analysis of variance (AOV) by ranks (NPARIWAY procedure; SAS 1990) was used to assess main effects. When a significant difference was detected, a Mann-Whitney U test (NPARIWAY procedure; SAS 1990) was used to locate those dif- ferences. Statistical significance for these and all other analyses was set at a= 0.05. Fish Sampling and Analyses Fish were collected during each sample period in the backwaters and from sandbar habitats with a bag seine (30-m long X 1.8-m high, 1.8-m* bag, 6-mm bar TABLE 1. Sampling schedule used in 1997 and 1999 in the study area, including the Missouri River main channel (CH), sandbar complexes (SB), and associated backwaters (BW). Habitats sampled during each sample period are denoted with an X. Some data, due to high flows and minimal habitat presence, were not available (NA). Habitats Sampled Sample Period Calender Days Month CH SB BW l 114-120 April X X X 2 139-145 May xX NA X 3 183-189 July X X X 4 225-231 August X X xX | 255-261 September X X X 28 THE CANADIAN FIELD-NATURALIST mesh). Six seine hauls were completed in each habitat type per sample period between 1000 and 1700 hours. The seine was pulled perpendicular to the backwater shorelines or with the current along sandbar margins for approximately 20 m and then arched to the shore- line. Captured fishes from the backwater samples were counted, measured to the nearest mm total length (TL), and released. Fishes captured during sandbar seining were also counted and measured; however, up to 30 Flathead Chubs from each of the following total length groups per sample period were euthanized and returned to the laboratory for biological analyses. The total length groups were < 60, 60-99, 100-139, 140-179, and > 179 mm. The Flathead Chub relative abundance was ex- pressed as number/seine haul and length-frequency histograms were created. Relative abundance data were tested for normality. Data not normally dis- tributed were transformed [log10 (n+1)] and again tested for normality. The relative abundance data were found to not be normally distributed, even after transformation; therefore, we used a nonparametric Mann-Whitney U test on the untransformed data to assess the differences between habitat types among sample periods within each year. The same analysis was used to evaluate differences between habitats during each sample period between years. To pro- vide additional samples for biological assessments in the two length groups exceeding 140-mm TL, a gill net (21-m long X 2-m high with one 7-m panel each of 1-, 2-, and 3-cm bar measure mesh) was drifted for short distances on the deeper portions (1.5-2.0 m) of the sandbar flats. Gill-net captured Flathead Chubs were added to samples for biological assessments; however, these data were not included in catch-per-unit-effort (CPUE) analyses. To better assess the potential utility of backwaters as nursery and rearing habitats for Flathead Chubs during their early life history, larval fishes were col- lected with modified quatrefoil light traps (Floyd et al. 1984; 25-cm high x 30-cm wide with two 2-mm and two 4-mm slot openings). Ten randomly placed light trap sets were completed in each backwater during each sample period. Attempts to sample lar- vae in calm channel areas with the light traps failed due to physical trap damage and sedimentation. The light traps were deployed between 1600 and 1800 h and emptied prior to 1100 h the following day. Two 12-h photochemical light sticks were used as the light source in each light trap each night. Photochemical light stick intensity and light duration varies with temperature; however, Kissick (1993) found that photochemical sticks attract larvae for at least 1 h. All photochemical sticks used in this study were found to continue glowing up to 24h after ini- tial use; regardless, light sticks were replaced each night. Larval fishes were preserved in 5% formalin and returned to the laboratory for identification. Case Vol. 116 specimens were sent to the Larval Fish Laboratory at Colorado State University for positive identification. Once identified, the specimens were enumerated and indexed as number/trap night. General Biological Evaluation In the laboratory, euthanized Flathead Chubs were again measured to the nearest mm TL and weighed to the nearest 0.1 g. Additionally, scales and otoliths were removed from September-sampled fish for age and growth analyses, stomachs were removed for a seasonal food habits assessment, and gonads of each fish were inspected to determine sex and maturity stage. Log10 transformed weight-length regressions were completed for each year and sample period (REG procedure; SAS 1990) and analyzed with an analysis of covariance (GLM procedure; SAS 1990) to determine if intercepts and slopes differed between years. Age and Growth Assessment We attempted to age the September-captured Flathead Chubs using both otoliths and scales. Scales were aged directly from a projected microfiche image. The focus, annuli, and radius data from each scale were digitized into the DISBCAL software program and mean length at age was determined using the Fraser-Lee method (Frie 1982). A regres- sion analysis of scale radius on total Flathead Chub length was conducted within DISBCAL to determine the best available intercept, or a-value, for the back- calculations. Otoliths were prepared for reading using both vertical cross-sectioning and horizontal grinding to help expose the otolith circuli; however, the annuli were not discernable. Therefore, all of the age and growth analyses were based on scale annuli. Mean length at age for Flathead Chubs from both years combined was determined to provide a general description of growth at this geographic location. To assess differences in growth between the high- flow year of 1997 and the mean-flow year of 1999, the mean growth increments for 2-cm TL Flathead Chub subgroups were determined for the first 258 days of each respective year. Martyn and Schmulbach (1978) suggested that annulus formation did not occur in Flathead Chubs until late May; therefore, the growth increment from the time of capture to the last annulus may only represent growth for 100—115 days. Using an AOV for unbalanced data sets, we complet- ed a two-way AOV with year and length category as the main effects (GLM procedure; SAS 1990). The interaction term between the main effects was not sta- tistically significant (P = 0.156); therefore, we com- pleted a one-way AOV between years within each length category to determine if the Flathead Chubs accumulated total length at a greater rate in 1997 or 1999. We also completed an AOV among length cate- gories within each year to determine if different length groups accumulated total length at significantly differ- ent rates. When a significant difference was detected, 2002 FISHER, WILLIS, OLSON, AND KRENTZ: FLATHEAD CHUBS IN UPPER MISSOURI 29 we used a Tukey multiple-range test to identify those differences (TUKEY option in GLM procedure; SAS 1990). Food Habits Assessment Stomach contents from the euthanized Flathead Chubs were enumerated. Food habits were assessed for feeding uniformity by determining the frequency of occurrence, while feeding intensity on each prey type was investigated by determining the percent by number for each prey item within each Flathead Chub length group. Frequency of occurrence was the pro- portion of individuals in each Flathead Chub length group that contained the prey item. The percent by number for each prey item was determined for stom- ach contents for each fish that contained food and the mean values for each length group were reported. Bowen (1996) noted that proportional diet data are not normally distributed; therefore, a Mann-Whitney U test was used to detect significant proportional differ- ences between years within each sample period and Flathead Chub length group. Because our objective was to assess food habit differences between different hydrologic conditions, no comparison among length groups within each year was completed. Results and Discussion Habitat Comparison Channel habitat parameters, including tempera- ture, dissolved oxygen, and turbidity were not signif- icantly different (P>0.05) from sandbar measure- ments in any collection period during either collection year (Table 2); therefore, the results and discussion will focus on the comparisons of backwa- ters with sandbar habitats only. Backwater and sand- bar temperatures were more similar in 1997 than in 1999. Backwater temperatures in 1997 were 0.4 to 1.4°C warmer than the sandbar habitats, but were not significantly different during any months (P > 0.05). In 1999, backwater temperatures were significantly higher (P< 0.05) in all months by 1.8 to 4.5°C (Table 2). September was the only sample period in 1997 with significantly higher backwater dissolved oxygen levels (P< 0.05); whereas in 1999, July was the only sample period without significantly greater backwater dissolved oxygen concentrations. Turbidity levels were significantly lower (P< 0.05) in the backwaters during all sample periods in 1997 and 1999 (Table 2). Sandbar habitat temperatures during the July and September sample periods of 1997 were significantly warmer (P< 0.05) than in 1999 by 1.6 to 1.7°C. In contrast, the backwaters were significantly cooler during May and September of 1997 (P< 0.05) than the same sample periods of 1999 by 4.1 and 2.4°C, respectively (Table 2). During July, August, and September of 1999 in both sandbar and backwater habitats, dissolved oxygen levels were significantly TABLE 2. Mean temperature, dissolved oxygen, and turbidity data collected in the Missouri River, North Dakota, in 1997 and 1999. Data were collected at designated time periods in backwaters (BW), the main channel (CH), and near sandbar complexes (SB). Standard errors are noted in parentheses. The results of three separate statistical comparisons are described in this table. Within each year and month, the three habitats were compared and the statistical results are coded with the letters a, b, and c. Within each year and habitat type, the sample periods (months) are compared and the statistical results are coded with the letters m, n, 0, and p. Within each habitat type and month, the 1997 and 1999 statistical compari- son is coded with the letters x and y. Within each set of comparisons, means with the same letters were not statistically dif- ferent (P>0.05). Each mean was derived from five measurements. 1997 Parameter/Period BW CH Temperature (°C) April amx] ]_7 amx] ] 2 May anx] 4.2 amnx ] 3.6 July a0x22.8 apy21.3 August aox2 1.9 aox19.9 September anx] 6,7 any16.0 Dissolved oxygen (mg/L) April anxQ 1 (0.3) aox8 5 (0.2) May any9 3 (0.3) apy9.6 (0.2) July amx7 () (0.2) amx6,7 (0.1) August amx7 7 (0.3) amnx7 2 (0.2) September amx8 3 (0.1) bnx7.7 (0.1) Turbidity (NTU) April ax66 (18) bnyQ4 (7) May amx]7 (8) bpy216 (24) July amx] 6 (3) boy 156 (6) August amx] 4 (5) bny85 (8) September amx]] (2) bmx38 (5) 1996 SB BW CH SB amx] 1.3 amx 12.6 bmx10).6 bmx1().8 NA any1$.3 bay 16.1 NA aoy2 1.4 a0x22.8 box] 9.4 box19.8 aox} 9.9 aox23.3 box19.8 box2().0 any 16,3 any]9.] bnx] 4.4 box 14.6 a0x8.6 (0.3) any9 4 (0.2) bax§.3 (0.1) bmx8 5 (0.2) NA amx7 8 (0.3) bmx§.6 (0.1) NA amx6.8 (0.2) amy8.4 (0.3) any8.5 (0.1) amy8.4 (0.2) amax7 .2 (0.3) any9 3 (0.2) bny8 3 (0.1) bmy§.1 (0.2) bax] 7 (0.1) ay} 1.0 (1.0) boy9 2 (0.1) bayQ 3 (0.2) bnyQ9 (5) anx46 (1 1) bnx7 4 (9) bax (3) NA anx33 (10) bex135 (2) NA boy 160 (6) amx } () (3) box] 12 (3) box] 16 (5) bny9 (3) amx§ (1) bnx65 (3) bnx7? (4) bmx39 (5) amxQ (2) bmx4() (3) bmx 39 (4) 30 THE CANADIAN FIELD-NATURALIST higher than in 1997 (P< 0.05; Table 2). Turbidity levels were not significantly (P> 0.05) different in the backwater habitats between 1997 and 1999, even though sandbar turbidity levels were significantly higher (P< 0.05) in April, July, and August of 1997 (Table 2). In 1997, the high flow created more floodplain connection points with longer duration. As a result, the 1997 temperature and dissolved oxygen levels in the backwaters were quite similar with those found in the channel. In 1999, these same variables operat- ed more independently with less influence from channel connections (Table 2). Regardless of year, the backwaters maintained the ability to filter chan- nel connection inflow and without exception, back- waters had significantly lower turbidity levels (P< 0.01; Table 2). This filtering capability, warmer water temperatures, an abundance of potential food resources such as zooplankton and benthic inverte- brates (Fisher 1999), and more static conditions would appear to create conditions sought and pre- ferred by some native fishes as nursery, rearing, feeding, and refuge habitats. Fish Sampling In 1997 and 1999 combined, 817 larval fish were captured in the light traps; however, none were veri- fied as Flathead Chubs. Of these larvae, 70% were Ictiobus spp. (Buffalo fishes), 12% were Common Carp, Cyprinus carpio, and 8% were River Carp- sucker, Carpiodes carpio. Additionally, only eight Flathead Chub juveniles and adults (>3-cm TL) were captured with the bag seine in the backwaters (Table 3). Significantly more Flathead Chubs, >12/seine haul, were captured in the sandbar habitats during all sample periods and both years (P< 0.01; Table 3). Although greater numbers of Flathead Chubs were captured on sandbar habitats, there also existed sev- eral significant differences in mean CPUE among sample periods. During both years, greater numbers of Flathead Chubs could be sampled near sandbar Vol. 116 - complexes later in the year (Table 3). In addition to higher CPUE, more large Flathead Chubs, particular- ly those exceeding 16-cm TL, could be captured dur- ing August and September, with the greatest range in size structure occurring in September during both 1997 and 1999 (Figure 2). Even though numerous larval fishes were sampled in the backwater habitats, no chubs of any genera were captured in the light traps. If the chub larvae were present, they were not sampled, possibly due to negative phototaxic habits or data collection timing problems. Likewise, the low CPUE indicated mini- mal presence of juvenile and adult Flathead Chubs in the backwater habitat. We hypothesized that high backwater connectivity to the channel would allow increased Flathead Chub use of the backwater habi- tats. Even in the presence of near record connectivity during 1997, Flathead Chub use of the backwaters as physical habitat was limited. The increased presence of large (> 16-cm TL) Flathead Chubs on the sandbars in the September samples (Figure 2) might suggest that the primary breeding adults stage, spawn, and recover in other channel habitats, as their conspicuous absence dur- ing the entire season except for the fall samples was easily noted. Peters and Holland (1994) found that all life stages of Flathead Chubs in the Platte River, Nebraska, utilized channel regions with sandy sub- strates and depths <50 cm. The sandbar habitats appeared to be important feeding and rearing areas for juvenile Flathead Chubs during all sample peri- ods. In addition, the age-0 Flathead Chubs, repre- sented by newly recruited individuals <4-cm TL, appeared on the sandbar habitats in late summer and early fall. The observation of age-0 Flathead Chub using the sandbar complexes could be an important life history documentation. Sexual Maturity Gonadal assessment in September-captured Flathead Chubs revealed that some males began to exhibit maturing testes by 60-mm TL, but likely did TABLE 3. Catch per unit effort for Flathead Chub captured with bag seines (number/seine haul) in two upper Missouri River backwaters (BW) and on two sandbar complexes (SB). The standard error of each mean is noted in parentheses. There are two sets of statistical comparisons denoted in this table. Within each habitat type and month, the letters a and b represent the comparisons between the 1997 and 1999 data points. Within each habitat and year (i.e., each column of means), the letters x and y represent the comparison among the five sample peri- ods (months). For each set of comparisons, means with the same letters are not statistically different (P > 0.05). Each mean was derived from six measurements. 1997 1999 Sample period BW SB BW SB April ax(),2 (0.1) bx12.3 (3.2) ax().2 (0.1) bx18.7 (8.2) May x0).2 (0.2) NA x0.3 (0.2) NA July ax(),3 (0.2) bxy24.2 (3.1) ax().2 (0.1) bx12.5 (6.3) August ax().0 (0.0) by37.0 (13.3) ax().() (0.0) by52.0 (11.5) September ax().0 (0.0) bxy29.2 (9.7) ax().Q (0.0) by55.8 (22.5) 2002 25 20 15 10 5 0 52 April Frequency 39 26 July Frequency 13 0 — 70 56 August Frequency 14 24 September Frequency 8 0 + S 2 7 20 a4 Total length (cm) FISHER, WILLIS, OLSON, AND KRENTZ: FLATHEAD CHUBS IN UPPER MISSOURI ie ‘a "aE a 30 24 18 12 220 165 110 55 0 70 56 42 28 16 20 a Bin tz 24 Total length (cm) FIGURE 2. Length-frequency histograms of Flathead Chub captured from sandbar habitats in the Missouri River, North Dakota during four sample periods in 1997 and 1999. The frequencies represent the total catch from six seine hauls per sample period. not spawn until the following season. Of male Flathead Chubs captured in April, 66% had devel- oped testes at 75-mm TL. At 110-mm TL, 100% of male Flathead Chubs had reached sexual maturity. The smallest Flathead Chub that contained devel- oped ovaries in April was 85-mm TL. Only 21% of the chubs that appeared to be female and were less than 125-mm TL contained mature eggs. By 155- mm TL, 80% of the female Flathead Chubs were mature, but not until 170-mm TL was 100% female sexual maturity noted. Olund and Cross (1961) suggested that sexual maturity was attained by both sexes at about 85-mm standard length in Alberta, and Brown (1971) noted well-developed gonads in Flathead Chubs approxi- mately 82-mm TL in Montana. Conversely, Gould (1985) found that the minimum total length for male and female maturation in the Musselshell River, 32 THE CANADIAN FIELD-NATURALIST Vol. 116 - TABLE 4. Frequency of occurrence (%) for stomach contents of Flathead Chubs collected in 1997 and 1999 from sandbar habitats in the Missouri River, North Dakota. Sample periods, total number (N), and percent of empty stomachs (E) are noted. Food items included Ostracoda (OST), Diptera (DIP), Ephemeroptera (EPH), Hemiptera (HEM), Hymenoptera (HYM), Odonata (ODO), Orthoptera (ORT), Plecoptera (PLE), Trichoptera (TRI), Copepoda (COP), Coleoptera (COL), and macrophytes (MAC). Total length groups (TL; mm), included 1=<60, 2=60-—99, 3=100—139, 4=140-179, and 5=>179. Absent groups imply that no specimens were collected. Month Year Length N B.S OST». DIP. BPH April 97 i 8. Oe LOO” 12.5" 6:0 April 99 1 13 OO wtGA- § 23:1°°""0:0 April 97 2 20 onn Sor, 26:7 ° O10 April 99 Z SOF 13s3 S35, AO) 35 April oF 3 13), 2O@), + 69.24. 20:8, 010 April 99 3 29) LAOS 6:9... 352°) t34 April 97 + 4 a3) Za0, 50.0. 0:0 April 99 4 10, = 0.0 00 20.0 200 July 97 1 af a ks mes 22 39 00 July 99 1 SU S00) GO 13.5 0:0 July 97 a 30°. 50:0") Sag" 13:3 6:0 July 2) 2 29) 138 0.0 69° 070 July oF, 3 10: 5 0:0), Fan@), S00 0:0 July ee) 3 S000 GO 16:7" 0.0 July oF 4 30:0) 66:6 0:07053.3 July 99 + 14. 14.3 0.0 0.0; 0.0 July o7 5 2» OR, 50:0 S08 YOO July 99 5) 3 338 0.0 0.0 0.0 August 97 1 9 OM 55:6, 33:3. “O10 August 99 1 l 0.0 0.0 100.0 9.0 August 97 2 30\ S333 333) 13.2 August 99 2 30 69 0.0 50.0 46.7 August 97 3 13: O05 3.8 Pini. August 99 3 2d op AOA 0.0 40.7 85.2 August 97 4 AN O05 75.0 0.0 50.0 August 99 = 30h vy. “ON 0.0 40.0 73.3 August 99 S 4 0.0 0.0 50.0 100.0 Sept oF 1 30° 167 50:0: 300° Aho Sept 212) 1 13° 30% OOO “iS:4", 010 Sept 97 2 30) Oa 56.7) 3323: (SAS Sept 28) 2 30 46.7 OO 16st . 010 Sept 97 3 20 siGes 70.0" 25.0. 530 Sept 99 5 28 60.7 0.0 Tek OO) Sept oo 4 I3.5feOies6.L 53.8 I4 Sept ie. 4 230 S45 OO N74 00 Sept All a 12 eels: Ss ALF 50.0 Sept 99 5 10 70.0 O07 16:0 0:0 Montana, was 113 and 123 mm, respectively. Al- though there is some disagreement about length at maturity, our results suggest that some males are reproductively active at age | and are all mature by age 2, whereas only a very few females are mature at age | and not until age 3 have all females entered the breeding population. Scarnecchia et al. (2000) also noted mature males at age | and a few mature females at age 2 in the Yellowstone River, Montana. Our observations are similar to Bishop (1975) and Martyn and Schmulbach (1978), who suggested that . sexual maturity was attained by age 2 and 105-mm standard length for the species as a whole. Food Habits In 1997, the most frequently consumed prey HYM ODO ORT PLE TRI COP COL IAG 2.0 00 00 00 00 125 S00" 360 ee O.0Ere 727 00° 0.0 00. 00 154° B27 20.0 00 633 . 00 | @0° 16:7 2267) ee 99 0.00.0" °0.0'" 13.3° S39 Sse 30.8. 0.0 308 00 00 53:3 335 0G ao 17.2. 17.2. O00 13.8. 206 \ 32 00 00, 25.0 250 00 0 3a 10.0 100 0.0 50.0 300 OG Bere 294 00 00 00 00 128 4°23 00 O00 OO 00 “0.0 (4383 3235257 20.0 00 0.00 00 (67 167) iw 00 00 00 00 7935 34° Rt Stee 30.0 00 0.0 50:0. 00° 100 300 a0 0.0 00 00 0:0... 6:7) 933) 523 tec 33.3... 0.0.....0.0,, 66.60 010) "66:GS3-2 ee 00 00 0.0. 00 . 00 78.6 \1As ees 50.0 0.0 00 1000 00,5 00 C0 it 00 00 00 00 00 333 232303 444 00 00 00 00° 333 33:3) 7a 00 00 00 00 §0:0,)0.0 OG Itc 19:3" 0.0 3.3 3.3 0.0°93:3° 250 eee 00 00 00 00 67 434° 33 Ga 53.8 0.0 385 462 00 308 30:59 Gita 00 00 00 00 00 111. 00 Sea 30.0.,.0.0 0.0. 50.0 » 0099300, (00a OO) pd 33 0.0) 6:7 30.0,.90:0 AO 0.0 00 00 00 500 500 (00. 46.7 0.0 00 00 00 2009300 Sata 0.0" 53.8 0.0 0.0 00 7.75) 00 isa ae 53.3 6.7 29.7 67 (3.3 19.8 902025 eee 0.0 43.3 0.0 00 00 00 (t= 232 —— 10.0 20.0 25.0 5.0 200 400 > 003 0.05 ¢ 17:9 0.0 0.0 0.0 10750 oat ae 30.8 00 00 461° 0.0 460° 0p eae Be Nas 2 | 00 00 00, 00> 00 174 2a 25.0) 98:3 0.0° 41.7 8.3 23.00 tese “iS ae 0.0 10.0 00 00 00, \00>) it tt 26 across all length groups of Flathead Chubs during the April and July samples were Ostracoda, Copepoda, Hemiptera (dominated by Corixidae), and Diptera. During the late summer and into autumn, Ostracoda, Hemiptera, and Copepoda con- tinued to demonstrate frequent presence in the stom- ach contents; however, Ephemeroptera and Trichoptera became more prevalent in the diets (Table 4). The 1999 frequency of occurrence data indicated that fewer Flathead Chubs were consum- ing Copepoda and Hemiptera. In addition, Ostracoda were nearly absent from the diets. Alternatively in 1999, Flathead Chub feeding strate- gies appeared to be more focused on Coleoptera in the spring and early summer and Hymenoptera and 2002 FISHER, WILLIS, OLSON, AND KRENTZ: FLATHEAD CHUBS IN UPPER MISSOURI 33 Ma 1997 [item G99 Ostracods 35 Diptera < 60 28 48 6 21 * * 36 2 24 14 -- a 12 * 7 0 0 SS 24 Hemiptera 32 Trichoptera °o 5 18 5 12 a. E 6 0 << 30 Copepoda 80 eae 64 5 be E* ke e re * 32 a 6 16 0 0 <60 60-99 99-139 140-179 Total length group (mm) <60 60-99 99-139 140-179 Total length group (mm) FIGURE 3. Numerical proportions (%) of primary prey taxa consumed by Flathead Chubs collected in April of 1997 and 1999 from the Missouri River, North Dakota. For each pair of bars (1997 and 1999), a statistical comparison was conducted. Those pairs with a % designation were significantly different (P<0.05). macrophytic plant seeds in the late summer and autumn (Table 4). Numerically, Ostracoda dominated the Flathead Chub diets in 1997, with Copepoda, Hemiptera, Trichoptera, and Diptera also contributing (Figures 3-6). In 1999, Ostracoda, Hemiptera, and Copepoda were significantly less abundant in the Flathead Chub diets (P< 0.05) during numerous sample peri- ods and among most length groups (Figures 3-6). The significant decline in several prey taxa was off- set by significant increases in the numeric propor- tions of Coleoptera, Diptera, Trichoptera, Ephemer- optera, and Hymenoptera during several seasons and for various length groups (all tests were P< 0.05; Figures 3-6). The food habits assessment provided evidence that the diets of Flathead Chubs in the high water year of 1997 were substantially different than those observed in the typical water year of 1999. Olund and Cross (1961) noted that Flathead Chubs col- lected in Alberta rivers had diets similar to those reported here, primarily consuming Corixidae, Hymenoptera, Diptera, and Coleoptera. In addition, Brown (1971) suggested that terrestrial insects were an important food resource during portions of the year. Cellot and Bournard (1987) noted that backwaters produce a number of invertebrates relatively uncom- mon in channel habitats. Some of those backwater taxa were the primary contents noted in our Flathead Chub diet analysis in 1997, such as Copepoda zoo- plankton and Hemiptera. Although Ostracoda can be abundant in the backwater habitats, their origin is not clear. Pennak (1989) noted that seed shrimps (Ostracoda) are well-equipped to thrive in lotic and lentic habitats. The Corixidae and Copepoda, howev- 34 THE CANADIAN FIELD-NATURALIST Vol. 116 Mm 1997 abel et Sele. Ostracods * 3 e. Ag — | ee 8. 24 2 ae 2 0 ye Hemiptera sS = Q = fe) a. fe) = oO S 59 Copepoda 6 39 8 26 * £43 aes 0) e) 9 fe) rae) = sor NI 9 er 9% Total length group (mm) Diptera 44 33 Ze Trichoptera AO on Coleoptera 32 24 16 8 A) grr o0-\™ 0-4 A192 Total length group (mm) FIGURE 4. Numerical proportions (%) of primary prey taxa consumed by Flathead Chubs collected in July of 1997 and 1999 from the Missouri River, North Dakota. For each pair of bars (1997 and 1999), a statistical comparison was conducted. Those pairs with a %& designation were significantly different (P < 0.05). er, both tend to be confined to lentic waters and exhibit low densities in lotic habitats (Merrit and Cummins 1984; Pennak 1989). Therefore, the evi- dence suggests that the high flows in 1997 likely contributed to the altered diets by transporting prey taxa from the floodplain to the channel and/or by washing in-channel food resources away from sand- bars at a greater rate than during a typical flow year. In 1999, the consumption of Coleoptera during April and extending into the other sample months was pronounced. Although reported as Coleoptera, nearly all of the beetles observed in the Flathead Chub stomachs belonged to the family Cicindelidae (tiger beetles). Borror and White (1970) noted that tiger beetles, where present, tended to be very abun- dant and inhabited open shorelines and beaches. Numerous tiger beetles were observed on the sand- bars, and Borror and White (1970) also suggested that the larvae tend to burrow into sandy areas dur- ing their juvenile stages. We can only speculate about the mechanisms that made tiger beetles avail- able to Flathead Chubs; however, during the spring and summer of 1999, many tiger beetles were observed in the drift and it is possible that as sand- bars shift, late-stage larvae and possibly adults are expunged from their burrows and become more sus- ceptible to fish predation. Alternatively, Flathead Chubs may actively forage into the substrates. Live Flathead Chubs held in the laboratory demonstrated digging behaviors while foraging, at times burying themselves past their pectoral fins. In 1997, the absence of tiger beetles from the diet was conspicu- ous. It is possible that the inundation of sandbar habitats, which occurred earlier than normal in 1997, 2002 FISHER, WILLIS, OLSON, AND KRENTZ: FLATHEAD CHUBS IN UPPER MISSOURI 35 Mmm 1997 fo} “4999 100 * Diptera Hemiptera se 30 ~— 80 c 24 = * 18 a 40 12 fe) ao 20 6 98 , Ree: Ephemeroptera Orthoptera % 24 E Pigg 3 ‘= fe] 2. fe) = ou S 24 Trichoptera 4 Copepoda £18 18 * 5 12 12 a. £6 6 * y 0) ~) ~~) 9 i 0 ~) ~~) 9 VA) go? 99°" An Al LO co? oa: io Total length group (mm) Total length group (mm) Ficure 5. Numerical proportions (%) of primary prey taxa consumed by Flathead Chubs collected in August of 1997 and 1999 from the Missouri River, North Dakota. For each pair of bars (1997 and 1999), a statistical comparison was conducted. Those pairs with a % designation were significantly different (P < 0.05). removed the staging substrates, either repelling the beetles away from the river bottom in search of bet- ter habitats or transporting the organisms out of the feeding area. The consumption of backwater-originated prey resources by Flathead Chubs during 1999, and more extensively in 1997, suggests that backwater prey production could be important to Flathead Chubs in the Missouri River, particularly during high-flow periods. In a relatively backwater-poor section of the Yellowstone River, Montana, upstream from our study site, Scarnecchia et al. (2000) also collected Flathead Chub in 1997, but found that 98% of the stomachs were empty. The high 1997 flow rates may have transported more typical Flathead Chub prey into unusable areas; therefore, the increased con- sumption of backwater-oriented food resources may have been a compensatory effort to obtain nutrition. We did not document any empty stomach propor- tions greater than 70%, and for most Flathead Chub length categories and sample periods, empty stom- ach proportions were typically < 40% (Table 4). It is unclear if the backwater prey availability was suffi- cient to provide adequate rations for the Flathead Chub population. Age and Growth To complete the back-calculation process, we needed to obtain a Flathead Chub intercept, or a- value, for the scale radius to total body length rela- tionship. A regression analysis based on all fish col- lected provided an a-value of 32 mm. This value seemed somewhat high considering that many small- er species, such as the darters, Etheostoma spp., typi- cally have a-values less than 22 mm (Carlander 1997) and larger species such as Walleye, Stizo- stedion vitreum and White Bass, Morone chrysops, 36 THE CANADIAN FIELD-NATURALIST Vol. 116 Mmm 1997 : Ea I"1998 Diptera Hemiptera se 30 30 S 24 24 ~ 18 18 & 12 12 2 ff 6 0 ia Hymenoptera 24 Orthoptera BS = 18 = 12 * =: * S 6 0 > 35 Trichoptera 25 Copepoda SS \e) = oF 15 x lk 8 14 10 * 2 so 9 Zoo goog Nt Total length group (mm) O 9 9 9 -) Ae) go? 09° 50°\" _\1 Total length group (mm) FIGURE 6. Numerical proportions (%) of primary prey taxa consumed by Flathead Chubs collected in September of 1997 and 1999 from the Missouri River, North Dakota. For each pair of bars (1997 and 1999), a statistical comparison was conducted. Those pairs with a *% designation were significantly different (P < 0.05). have a-values greater than 30 mm (Carlander 1982; Beck et al. 1997). To help validate the use of a 32- mm intercept during the back-calculations, we fur- ther assessed the Flathead Chub specimens in our collection that were <40-mm TL and in good body condition. Upon further inspection, all of the Flathead Chubs < 30-mm TL were scaleless and did not appear to be physically damaged in any way to indicate that scales may have been dislodged during handling. Of the remaining Flathead Chubs < 40-mm TL, 45% of those between 30- and 35-mm TL were scaled and by 38-mm TL, all specimens had scales with discernable circuli. Therefore, the a-value of 32 mm was used with relative confidence. Aging the Flathead Chub scales was a difficult pro- cess and the agreement proportion between the two initial readers (both inexperienced) was 42%. Likewise, Scarnecchia et al. (2000) indicated that Flathead Chub scale aging was challenging. Gould (1985) relied upon length-frequency histograms to identify year classes after unsuccessful attempts to age Flathead Chubs with scales, vertebrae, and opercula. The length-frequency histograms that we formulated for this Missouri River population did not allow us to confidently identify cohorts (Figure 2). Although early age groups can be discerned on the length-fre- quency histograms, variability was too high to rely on that technique. Therefore, we consulted other cyprinid researchers and aging experts. After receiving a cyprinid aging tutorial, we re-aged the scales with three readers (2 inexperienced and 1 experienced) and reached a consensus age on 96% of the scales. The remaining 4% were eliminated from the analysis due to our inability to assign an age. Our age analysis indi- cated that the maximum age of Flathead Chub in our collection was 5 years. Therefore, considering growth and maturation patterns, many females may only release eggs in one or two seasons. 2002 FISHER, WILLIS, OLSON, AND KRENTZ: FLATHEAD CHUBS IN UPPER MISSOURI 57 TABLE 5. Mean back-calculated total lengths (TL; mm) at age for Flathead Chubs collected in 1997 and 1999 from the Missouri (MO) River in North Dakota. Standard errors are in parentheses. Also noted are total lengths (mm) at age for Flathead Chubs from Perry Creek, Iowa (Martyn and Schmulbach 1978) and from the Peace River, Alberta [Bishop (1975); fork lengths converted to total length using conversion in Martyn and Schmulbach (1978)]. Age MO River (N) MO River (TL) Perry Creek Peace River 1 85 104.1 (2.8) 77.8 (0.6) 100.4 (NA) 2 a7 152.9 (4.7) 109.6 (0.5) 123.6 (NA) 3 17 186.4 (3.8) 130.0 (0.5) 152.2 (NA) 4 6 223.2 (6.9) 148.0 (2.3) 164.9 (NA) 5 1 267.0 (NA) NA 179.2 (NA) The back-calculated lengths at the time of annulus Incremental growth analyses for Flathead Chubs col- formation for each age group suggested that Flathead lected in September of each year indicated that por- Chubs, on average, increase their length by approxi- _ tions of the Flathead Chub length range, including mately 46% between ages 1 and 2, but then growth _ those between the total lengths of 95 and 135 mm, slows and plateaus at annual length increases of significantly increased at a more rapid rate during 19-22% during the remainder of their life (Table 5). 1997 than did similar sized fish in 1999 (P< 0.02; Mean growth increment (mm) 50 =@— 1997 == 1999 45 40 35 30 25 20 165 gst ys yo 36" , \1 539% 49° Total length group (mm) FiGURE 7. Mean growth increments for Flathead Chubs collected in the Missouri River, North Dakota in September of 1997 and 1999. The increments represent estimated growth from the time of annulus formation to the time of capture each year. The letters a and b represent the statistical comparison between years for each total length group noted on the x-axis (vertical comparison). The letters x, y, and z denote the comparison results among total length groups within each sample year (horizontal comparison). Means with the same letters were not statistically different (P > 0.05). 38 THE CANADIAN FIELD-NATURALIST Figure 7). With the exception of those age-1 chubs less than 95-mm TL, all length groups had higher mean increments in 1997 than in 1999; however, dif- ferences for groups >134-mm TL were not statisti- cally significant (P< 0.05). In 1997, the longest growth increments were attained by chubs between 95 and 174 mm, whereas in 1999, chubs between 95 and 135 mm had significantly lower growth incre- ments (P < 0.05; Figure 7). As compared with the maximum age of 5 reported here, Martyn and Schmulbach (1978) found Flathead Chubs up to age 4 in Perry Creek, Iowa; however, Scarnecchia et al. (2000) suggested that Flathead Chubs in the Yellowstone River, Montana, reached age 7 and Bishop (1975) noted that the maximum life span of Flathead Chubs in the Peace River, Canada was 10 years. Additionally, the longest Flathead Chub in our sample was 275-mm TL, whereas Kristensen (1980) reported capturing 293 Flathead Chubs in the Peace-Athabasca River delta in Canada that had a mean TL of 345 mm. The sub- stantially higher total length and longevity reported for Flathead Chubs in the northern portions of the species range may be attributed to less disruption of recruitment patterns and cooler water temperatures that moderate metabolic activity, ultimately extend- ing the average potential life span. The population of Flathead Chub utilizing the sandbar habitats in this study was dominated by individuals < age 3. Brown (1971) reported lengths at ages 1-3 for Flathead Chubs in Montana; however, no indication was given pertaining to how these lengths and ages were derived or if the measurements were in total, fork, or standard length (Table 5). Martyn and Schmulbach (1978) aged Flathead Chub scales col- lected from specimens in Perry Creek, a Missouri River tributary in Iowa, and noted lengths consider- ably shorter for each age group than those reported in our study (Table 5). Gould (1985) also noted three Flathead Chub total length modes at 43-, 81-, and 116-mm TL in the March length-frequency his- togram from the Musselshell River, Montana, but did not suggest that the modes represented age groups. Our assigned ages and back-calculated lengths at age differed from several other reports, but were similar to those reported at ages 1—5 in the Vol. 116 - Peace River, Alberta (Bishop 1975). We believe that our aging was done appropriately and given the wide variability of habitats and phenotypical expression across the species range, our results should be repre- sentative of the Flathead Chub population found in the study area. Although understanding mean growth patterns for Flathead Chubs as a population is valuable, we were particularly interested in the growth rates during 1997 and 1999 when the diets significantly differed. The cohorts of Flathead Chubs in 1997 utilized prey that were likely backwater originated, including Copepoda and Hemiptera, at a significantly higher rate than the same cohorts did in 1999 (Figures 3-6). Food resources that appeared to have backwater origination were nearly absent from the chub diets in 1999. This dietary absence may have influenced the growth rates of different length groups to varying degrees, especially when considering feeding effi- ciency and food item assimilation patterns that can cause substantial changes in the bioenergetics of dif- ferent cohorts within a species (Adams and Breck 1990). In addition to changes in diet, environmental con- ditions can also alter bioenergetic functions and affect fish growth rates and body condition by directly influencing metabolism and prey dynamics (e.g., Kitchell et al. 1977; Swenson 1977). In 1997, water temperatures were cooler in the early portions of the annual flood pulse (i.e., April and May), but were significantly warmer (P< 0.05) from July through September (Table 2). The higher summer and fall temperatures in 1997 may be attributable to the extended exchange of water between the flood- plain and the channel. Waters retained in backwaters and other floodplain wetlands tend to be substantial- ly warmer, as was demonstrated in 1999 when the backwaters functioned in an uncoupled manner from the channel (Table 2). The higher temperatures in 1997 may have contributed to the increases in growth increments that could also be noted on the scales from fish collected in 1999, particularly those that were ages | and 2 in 1997. Body Condition Regardless of sample period, Flathead Chubs cap- tured in the mean flow year of 1999 weighed less at TABLE 6. Weight-length equations for Flathead Chubs captured in various sample periods in 1997 and 1999 from the Missouri River in North Dakota. Statistical comparisons (analyses of covariance), represented by their p-values, between years for intercepts (a) and slopes (b) are noted. Weight (wt) is in grams and total length (TL) is in mm. Weight-length equations Period 1997 April log ,pwt = -5.369+3.065(log ,, TL) July log ,pwt= -5.621+3.226(log,, TL) August log wt = -4.9024+2.853(log,, TL) September log ,ywt = -5.349+3.063(log, TL) P-values 1999 a b log ,)wt = -4.680+2.802(log,,TL) 0.001 0.001 log wt = -5.028+2.997(log,, TL) 0.001 0.001 log ywt = -5.2454+3.107(log,,TL) 0.006 0.001 log, )wt = -4.918+2.938(log,, TL) 0.001 0.001 2002 FISHER, WILLIS, OLSON, AND KRENTZ: FLATHEAD CHUBS IN UPPER MISSOURI 39 + 1997 @ 1999 log, Weight (g) oS ee eS mp) Nn =) Nn mn) S Nn a lididicn AB’ Ain 2D, ».221 Ls — August log, Weight (g) SP re sa Nn =) Nn S a) ened rome log; ototal length (mm) pe | et | 22 Ai ythetaiA By 39>. 20 ite nd deed 4 + September EE 2M Beleth AeKGiy {bee 9 42s (1d De oti, oath log, total length (mm) FiGuRE 8. Weight-length relationships (log10 transformed) from Flathead Chubs collected in the Missouri River, North Dakota, during various periods of 1997 and 1999. The cross-hair markers denote chubs collected in 1997 and the circles represent those collected in 1999. The regression relationship for each set of data is depicted with a solid line. Statistical analyses for these data are summarized in Table 6. all comparable lengths than in 1997 (Figure 8). The intercept values and slopes of these regression analy- ses were all significantly different between years during each sample period (P< 0.006; Table 6). During some sample periods, the 1999 predicted weights for some Flathead Chub lengths exceeded the 1997 predicted weights by as much as 50%. The weight-length equations reported here were similar to those documented by Gould (1985); how- ever, that study did not identify any significant dif- ferences in slopes or intercepts among sample peri- ods. It would be reasonable to hypothesize that when the environment favors increased growth rates (based on total length), that the habitats would also be favorable for individuals to maintain high body condition (plumpness). Although the logic of this argument seems apparent, our data suggest that Flathead Chubs in 1997 acquired greater total length additions, but did so while they were less plump. Fish condition has been correlated with prey avail- ability in many lentic populations (Wege and Anderson 1978; Liao et al. 1995; Marwitz and Hubert 1997; and Porath and Peters 1997), but we are aware of no such analyses for lotic fish popula- tions. We cannot explain the contradictory condition and growth observations discussed above; however, it could be hypothesized that during a year when flow velocity is high (e.g., 1997) it is ecologically advan- tageous for a fish to be more fusiform than when under conditions of lower flow rates. Fisher et al. (1996) noted that healthy burbot, Lota lota, popula- —— aes. nee 6 eee eee 40 THE CANADIAN FIELD-NATURALIST tions from lotic systems consistently maintained lower overall body condition; however, annual varia- tion in these condition measures was not assessed. Although Flathead Chubs in 1997 were thinner and had to cope with higher turbidity levels (Table 2), the population was also provided with food resources and warmer water temperatures due to the higher rate of floodplain interaction. Scott and Crossman (1973) noted considerable morphological variation in Flathead Chubs across its North American range and Olund and Cross (1961) sug- gested that subspecies, based on morphological fea- tures that included differences in total length to cir- cumference ratios, be recognized. However, Bailey and Allum (1962) hypothesized that the manifesta- tions were strictly related to environmental condi- tions, not genetic strains. The substantial differences that we observed between two years within the same Flathead Chub population support the plasticity sug- gestion of Bailey and Allum (1962). Flathead Chub food habits are often referred to in general, but seldom supported with documentation (e.g., McLane 1978; Baxter and Stone 1995). Brown (1971) reported that biologists in Montana believe the Flathead Chubs feed on aquatic invertebrates, extensively utilize abundant terrestrial insects that become part of the drift, and are sufficiently omniv- orous to feed on vegetative matter. Our results sup- port these assumptions, including the sporadic feed- ing on plant tissues. Our data indicate that diets vary substantially during differential hydrograph cycles and, when combined with varied flows and other environmental conditions, influence growth rates and condition. The backwater-produced prey appeared to be utilized by small to mid-sized Flathead Chubs. We were unable to determine if Flathead Chubs prefer the backwater organisms or are using their availability to compensate for absent channel resources during high flow periods. Riverine ecologists (e.g., Kennedy 1979; Eckblad et al. 1984) have assumed that backwater habitats are critical to native fishes evolved in large river sys- tems. We documented limited direct physical use of backwater habitats by Flathead Chubs during any life history stage; however, the indirect benefits of floodplain prey production and water conditioning are possibly critical during high flow hydrographs. Acknowledgments We thank the U.S. Fish and Wildlife Service (Region 6), the South Dakota Cooperative Fish and Wildlife Research Unit, South Dakota State Uni- versity, and the U.S. Geological Survey Species at Risk program for funding and administering this research. We also express our appreciation to Charles Pyle, Randy Sheik, Steve Wilson, Nate Olson, Dan Moon, Ryan Doorenbos, Danielle Johnson, and Gene Galinat for their assistance with field data collection and laboratory analyses. We Vol. 116 - thank Mike Brown for statistical advice, several reviewers, and the North Dakota State Department of Fish and Game, particularly Fred Ryckman and Greg Power for their cooperation and logistical assistance. This manuscript has been approved for publication by the South Dakota Agricultural Experiment Station as Journal Series Number 3187. Documents Cited (marked * in text) USGS (U.S. Geological Survey). 1999. Water resources of Montana: historical water data files. http://waterdata. usgs.gov/nwis-w/mt. Literature Cited Adams, S. M, and J. E. Breck. 1990. Bioenergetics. Pages 389-415 in Methods for fish biology. Edited by C. B. Schreck and P. B. Moyle. American Fisheries Society, Bethesda, Maryland. Amoros, C. 1991. 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Received | May 2000 Accepted 5 March 2002 Abundance and Distribution of Breeding Waterfowl in the Great Clay Belt of Northern Ontario R. KENYON Ross!, KENNETH F. ABRAHAM2, TED R. GADAWSKIF, ROBERT S. REMPEL4, T. SHANE GABORS, and RON MAHER® ‘Environment Canada, Canadian Wildlife Service, Ontario Region, 49 Camelot Drive, Ottawa, Ontario K1A 0H3 Canada [Corresponding Author] Ontario Ministry of Natural Resources, Science, Development and Transfer Branch, 300 Water Street, 3" Floor N, Peterborough, Ontario K9J 8M5 Canada 5R.R. # 2, Shanty Bay, Ontario LOL 2L0 Canada ‘Ontario Ministry of Natural Resources, Centre for Northern Forest Ecosystems Research, Lakehead University Campus, Thunder Bay, Ontario P7B 5E1 Canada ‘Institute for Wetland and Waterfowl Research, c/o Ducks Unlimited Canada, Oak Hammock Marsh Conservation Centre, P.O. Box 1160, Stonewall, Manitoba ROC 2Z0 Canada Ducks Unlimited Canada, 614 Norris Court, Unit # 1, Kingston, Ontario K7P 2R9 Canada Ross, R. Kenyon, Kenneth F. Abraham, Ted R. Gadawski, Robert S. Rempel, T. Shane Gabor, and Ron Maher. 2002. Abundance and distribution of breeding Baer! in the Great Clay Belt of northern Ontario. Canadian Field- Naturalist 116(1): 42-50. The abundance and distribution of breeding waterfowl in the Great Clay Belt of northern Ontario was determined through helicopter surveys of 117 fixed plots (2 X2 km each) during the nest initiation periods from 1988 to 1990. This area has higher fertility, flat topography, high water table and better access than the surrounding Boreal Forest, and therefore has greater potential for increased waterfowl production through habitat management. Overall breeding density averaged 112.5 indicated breeding pairs per 100 km?, 68% being of the four most common species [Mallard (Anas platyrhynchos), Ring- necked Duck (Aythya collaris), American Black Duck (Anas rubripes), and Common Goldeneye (Bucephala clangula)); 13 other species were encountered. The average total of breeding waterfowl for the region was estimated at 59330 pairs. Distributions of the species were related to ecodistrict and to surficial geology. The more northerly of the two main ecodis- tricts had higher densities of American Black Ducks, Ring-necked Ducks, Common Goldeneyes, and Canada Geese (Branta canadensis). Mallard and Hooded Merganser (Lophodytes cucullatus) distributions correlated with presence of sur- ficial clay and moraines, respectively. Less common species including Green-winged Teal (Anas crecca) and American Wigeon (Anas americana) appeared to be concentrated in smaller-scaled habitat features (beaver pond sequences and estu- arine marshes, respectively). Results generally agreed with those of earlier Clay Belt surveys. Total breeding density of waterfowl is slightly higher than that of surrounding regions. Key Words: ducks, populations, habitat, boreal, forest, Ontario The boreal forest region in eastern Canada has methods were applied that breeding density esti- been largely ignored as an important area for water- mates for the full range of waterfowl species could fowl production owing to its relatively low fertility be made (Dennis 1974; Ross 1987). These intensive and often difficult accessibility when compared to _ surveys revealed that different physiographic units of the prairies. An initial assessment by Hanson et al. _ the boreal forest had characteristic waterfowl species (1949) based on the limited information available at compositions. One such unit is the Great Clay Belt, the time concluded that the area was of minimal also known as the Northern Clay Section (B4 in importance to most waterfowl populations. How- Rowe 1972), and hereafter termed the Clay Belt in ever, aS more quantitative surveys were applied, ini- this paper. In Ontario, the Clay Belt extends east tially by the U.S. Fish and Wildlife Service in the from Hearst to the Quebec border and south approxi- 1950s and 1960s (Addy et al. 1952, Kaczynski and mately to Timmins (see Figure 1); the remainder of Chamberlain 1968), it became clear that, while this unit occupies an area of roughly similar size in breeding densities were low, the vast area of the contiguous Quebec. The Clay Belt is unique in hav- boreal forest made its contribution to continental ing a rich clay soil in contrast to low fertility habitats waterfowl stocks substantial (Wellein and Lumsden of muskeg and exposed-bedrock shield surrounding 1964). The accuracy of these earlier surveys was _ it. The higher fertility of the Clay Belt is evidenced limited by the difficulty of observing waterfowl in by higher rates of wetland occupancy by waterfowl forested wetlands from fixed-wing aircraft . than in adjoining areas (Dennis 1974). Moreover, the (Chamberlain and Kaczynski 1965). In Ontario, it combination of its general fertility and flat topogra- was only when more intensive and efficient survey phy, high water table, and relative accessibility due 42 2002 FEeolo NE BAe Ross, ABRAHAM, GADAWSKI, REMPEL, GABOR, AND MAHER: BREEDING WATERFOWL 43 Human Geography ical Classification Lac Matagami Ecoregion Ecodistrict, 29 Ecodistrict,.30 Other Ecodistricts James Plain Ecoregion Chapleau Plains Ecoregion Surficial Geology Lacustrine Sediments (Clay) Ground Moraine End Moraine Marine Sediments FIGURE 1. Geographical locations, ecological classification, and surficial geology units of the Great Clay Belt of Ontario. to the extensive network of roads for logging and mining, gives the Clay Belt greater potential for enhanced waterfowl production through habitat man- agement than in the neighbouring boreal forest. This importance is well-recognized and the Clay Belt is included as a priority area in the Eastern Habitat Joint Venture of the North American Waterfowl Management Plan (NAWMP Update 1994). Maximizing the effectiveness of management activities requires knowledge of the breeding distri- butions and habitat preferences of various waterfowl species frequenting the area. A study of waterfowl productivity across the Clay Belt of Ontario was therefore undertaken jointly by Ducks Unlimited Canada, the Ontario Ministry of Natural Resources and the Canadian Wildlife Service. The main objec- tives were to: (1) assess species composition, densi- ty, and distribution of the waterfowl community, (2) determine habitat correlates of nesting and brood- rearing waterfowl, and (3) develop a wildlife habitat map using LANDSAT. This paper describes water- fowl species composition, density and distribution across the Clay Belt, relates these to broad habitat patterns in the area, and estimates the contribution of the Clay Belt to overall waterfowl populations in northern Ontario. 44 THE CANADIAN FIELD-NATURALIST Methods Study Area The Clay Belt is a relatively flat, humid forest region corresponding to an area occupied by former glacial lakes Ojibway and Barlow (Rowe 1972; Sharpe and Brodie 1931). Bedrock is mostly overlain by sedimentary clays and extensive peatland. There are relatively few large lakes, and streams and rivers are generally low gradient compared to adjacent exposed Precambrian shield areas. Tree species of the Clay Belt reflect this topography and geological history. The main tree species is Black Spruce (Picea mariana) which occurs on flat lowlands and moister uplands. In areas of better local drainage, species include White Spruce (P. glauca), Trembling Aspen and Balsam Poplar (Populus tremuloides, P. balsamifera), White Birch (Betula papyrifera), and Balsam Fir (Abies balsamea). Jack. Pine (Pinus banksiana) occurs widely on sandy, drier sites such as eskers, moraines and outwash plains. Clay areas were cleared for agriculture in the 1930s and 1940s; these are limited in extent and distribution around communities on or very near Highway 11, which bisects the study area from southeast to northwest (Figure 1). Wetlands vary widely in size, and beaver impoundments on the many low gradient streams account for many of them. A wetland classification system for the area and an assessment of its rele- vance to duck species in the Clay Belt was devel- oped by Rempel et al. (1997) as part of this study. Population Estimation Distribution and abundance of breeding waterfowl in the Clay Belt was determined by aerial surveys of fixed plots during nest initiation periods in 1988, 1989 and 1990. These plots were selected using a two-staged process in which the whole Clay Belt was divided into a grid of 10 x 10 km blocks follow- ing the UTM system. Each block was then subdivid- ed into 25 —-2 X2 km plots, and one plot selected at random from each block for a total of 117 plots. This sampling regime allowed us to describe species dis- tribution, and to statistically estimate population size and habitat correlations. Surveys were carried out by helicopter (after Ross 1985, 1987). All surveys were done from a Bell 206B helicopter on high skids and equipped with a range extender on the fuel tank and bubble observa- tion windows on the back doors for improved visibil- ity. Two observers in the back watched for water- fowl from their respective sides of the aircraft, and passed observations by intercom to the navigator/ recorder in front. That person wrote these directly onto acetate-covered aerial photographs, recording species, sex, numbers, and exact locations of all waterfowl; care was taken to ensure that sightings came from within the boundaries of the plots. The aircraft passed over all suitable habitat in each plot at altitudes as low as 20 m, and at speeds ranging from Vol. 116 - a hover to 100 km/h. Multiple passes were made over those waterbodies where the initial coverage was thought to be inadequate or where birds seen were not initially identified. Because most wetlands in the Clay Belt are small and clearly bounded, total coverage of all wetlands in the plots was accom- plished by following shorelines, and no transects were employed. All plots were surveyed during the following periods: 10-21 May 1988, 20-26 May 1989, and 20-22 May 1990. Results of the survey are expressed as numbers of indicated breeding pairs (IBP) per species. For clear- ly dimorphic species, IBP was based on the total of lone males, pairs, and number of males in flocks with up to 5 males (as in Dzubin 1969). For Amer- ican Black Ducks (Anas rubripes) which are more weakly dimorphic, determination of sex was suc- cessful in about 90% of the cases; the remaining Black Ducks were ascribed indicated pair values based on the average proportions of indicated pairs to number of individuals in the three sighting classes (lone birds, two-bird groups, flock of three or more) as taken from all breeding pair surveys in northeast- ern Ontario from 1988 to 1990 (Ross, unpublished data). Indicated pairs for Canada Geese (Branta canadensis) were counted for each single, pair, or group of three. Counts of indicated pairs on the plots based on this helicopter survey technique are compa- rable to intensive ground survey results and are con- sidered sufficiently close to the real values to be used directly in breeding density calculations (Ross 1985); no visibility correction factors have been applied. Habitat Parameters Although the Clay Belt is designated as a single forest section by Rowe (1972), other classifications can be used to subdivide this area to examine broad patterns in waterfowl distribution. In Figure 1, we present two classifications of the region. The first is an ecological classification showing the boundaries of the ecoregions and ecodistricts (Wicken 1986; Wickware and Rubec 1989); an ecodistrict is defined as a part of an eco-region characterized by a distinc- tive pattern of relief, geology, geomorphology, vege- tation, soils, water, and fauna. A second classifica- tion portrays substrate and is based on the surficial geology of the region (Glacial Map of Canada, Geo- logical Survey of Canada). To aid in interpretation of waterfowl distribution data, amounts of various potential nesting habitats (wetland associated with standing water) in each plot were measured by elec- tronic planimeter, and number of water bodies were counted. Statistical Analysis Overall breeding densities in both ecodistrict and substrate units were compared using a Mann- Whitney U Test (from Siegel 1956). Contour maps 2002 Ross, ABRAHAM, GADAWSKI, REMPEL, GABOR, AND MAHER: BREEDING WATERFOWL 45 TABLE |. Breeding densities of waterfowl of the Clay Belt (present and earlier studies) plus estimates of population sizes from present study. Dennis Ross Source (1974) (1987) Present Study Mean (1988— Year 1973 1982 1988 1989 1990 1990) Parameters Breeding Density (IBP/100 km?) Population Size (range) Species Canada Goose — 1.0 1.98 3.24 2.36 2.52 1330 (1040-1710) Wood Duck - f.Q5: 0.54 2.88 BAF 1.86 980 (290-1520) Green-winged Teal oF 4.0 4.50 8.27 11.96 8.21 4330 (2370-6310) American Black Duck a3 10.2 16.09 19.85 20.09 18.72 9870 (8490-10600) Mallard 252 21.0 2A 22, 29.50 24.09 24.88 13120 (11190-15560) Blue-winged Teal 6.2 6.0 3.96 7.01 6.70 5.88 3100 (2090-3700) American Wigeon 17.8 — 2.16 2.88 J 3.00 1580 (1140-2100) Other Dabblers = - = 0.54 0.18 0.24 130 (0-290) Ring-necked Duck 12.4 18.0 16.01 22.66 24.09 20.90 11020 (8440-12710) Lesser Scaup - ~ 0.72 0.72 2.90 1.44 760 (380-1530) Common Goldeneye 27.8 11.0 11.51 11.70 13.22 12.41 6390 (6070-6970) Bufflehead — - 0.90 0.90 0.72 0.84 440 (380-470) Hooded Merganser P| 4.0 6.65 5.40 8.15 6.71 3540 (2850-4300) Common Merganser ~ 6.0 5.58 3.78 5.25 4.95 2610 (1990-2940) Other Divers 1.3 - - 0.54 0.18 0.24 130 (0-290) Total 132.8 82.2 91.81 119.85 126.07 112.50 59330 (48420-66490) of breeding pair densities and other parameters were generated using surfaces created by the potential mapping routine (POTMAP) of the SPANS geo- graphical information system (INTERA TYDAC Inc. 1991) and are based on sequential averaging of point values encompassed within an outer circle of 47 km of the each specific plot value; an inner circle of 10 km holds the specific plot value constant over that area. Results and Discussion Waterfowl Population Sizes Breeding densities of the more common waterfowl species for each of the three years are presented in Table 1. Densities, for each species, were similar in 1989 and 1990, but those for 1988 were consistently lower for dabbling ducks and Ring-necked Ducks (Aythya collaris). These differences reflected outlier values generated when very low counts were made in 1988 on a small number of plots that had much higher numbers in later years. Because annual varia- tion in waterfowl numbers in boreal systems is not well known and the ranges of values were less than 20 percent of the average for the most common species, we used three-year means to represent aver- age breeding densities of waterfowl species in the Clay Belt (Table 1). Overall breeding density of waterfowl was 112.5 IBP/100 km2; the four most common species (68% of total IBP) were American Black Duck, Mallard (Anas platyrhynchos), Ring-necked Duck, and Common Goldeneye (Bucephala clangula). Less common, but widely encountered (4 species, 23% of total IBP), were Green-winged Teal (Anas crecca), Blue-winged Teal (Anas discors), Hooded Mer- ganser (Lophodytes cucullatus), and Common Merganser (Mergus merganser). The remaining nine percent included five uncommon and locally occur- ring species [Canada Goose, Wood Duck (Aix spon- sa), American Wigeon (Anas americana), Lesser Scaup (Aythya affinis), Bufflehead (Bucephala albe- ola)], plus four rare species [Northern Pintail (Anas acuta), Northern Shoveler (Anas clypeata), Surf Scoter (Melanitta perspicillata), and White-winged Scoter (Melanitta fusca)]|. Estimates of breeding populations of all but the rare species (also in Table 1) were generated by multiplying average densities by the area of the Ontario Clay Belt (52 740 km2), to give an average total of 59 330 breeding pairs of waterfowl in this area (range: 48 420-66 490). Habitat Characteristics Only the two largest ecodistricts (Numbers 29 and 30) had sufficient numbers of survey plots (81 and 20, respectively; 86% of all plots) for separate analy- sis. Ecodistrict 30 had a lower elevation and a greater proportion of standing water than did Ecodistrict 29 (Wickware and Rubec 1989). Amount of waterfowl breeding habitat and number of water bodies per plot were significantly higher in Eco- district 30 (Table 2). Ecodistrict 29 contained most of the agricultural areas and much of the area that has been logged (Hutton and Black 1975). 46 THE CANADIAN FIELD-NATURALIST Surficial geology is another way of subdividing the Clay Belt (Figure 1). Although lacustrine sedi- ments, essentially clay, are the dominant substrate, much of the study area, particularly toward the west, is underlain by ground moraines with several smaller sections of end moraines. As clay usually contributes to greater ecological productivity, we expected that its presence should influence the distribution of some species. No statistical difference was found in number of water bodies or the total area of wetlands in plots of the two substrates (Table 2). Waterfowl Distribution Distributions of individual species are displayed as contoured maps based on mean breeding density per plot (Figure 2) for all species that were recorded on more than nine different plots. In Table 3, breeding densities of these species are compared between the two ecodistricts and between the two substrate types. The most abundant species was the Mallard that was found throughout the area with highest densities following a diagonal band (northwest-southeast) across the Clay Belt. There was no difference in breeding density between ecodistricts, but density was higher on plots with a clay substrate. Although habitat fertility influences Mallard distribution (Merendino and Ankney 1994), the area of highest breeding density is also that with most agricultural and tree-harvesting activities, which produce prairie- and parkland-like habitats to which Mallards are well-adapted (Bellrose 1980). American Black Ducks were abundant and widespread in the Clay Belt (second amongst dab- bling ducks, third overall). Highest breeding densities occurred in the north. Of the two ecodistricts, signifi- cantly higher breeding density was encountered in Ecodistrict 30, but unlike Mallards, no relationship was found between breeding density and substrate. Although Mallards and Black Duck are closely relat- ed and interbreed, their distributions did not show the same patterns even though they were among the most common species (see also Ross and Fillman 1990). Vol. 116 - The most common diving ducks in the Clay Belt (and second most abundant duck overall) were Ring- necked Ducks. Their density distribution was similar to that of Black Ducks, with which they regularly co- occur (McNicol et al. 1987). Ecodistrict 30 support- ed higher breeding densities, but there was no rela- tionship with substrate. Common Goldeneye was the fourth most abundant species, and like Black Ducks and Ring-necked Ducks, they occurred in significantly higher density in Ecodistrict 30 than in Ecodistrict 29, and there was no correlation between distribution and substrate. Green-winged Teal were observed throughout the study area but did not show a pattern explainable by either ecodistrict or substrate classifications, suggest- ing that the birds were using specific habitat types with distributions unrelated to these factors. In the Clay Belt, occurrence of this species is significantly associated with Beaver pond systems (Rempel et al. 1997). Paquette and Ankney (1996) demonstrated high specificity in wetland use by this species in British Columbia. Hooded Mergansers occurred in moderate num- bers. High density zones largely coincided with those of Common Goldeneye, which may reflect the distribution of suitable nesting cavities for which the two species have similar requirements (Lumsden et al. 1980). No relationship with ecodistrict was evi- dent, but breeding density was significantly higher in areas with moraine substrate than in those underlain by clay; the latter substrate may produce greater tur- bidity in waterbodies making them less suitable for foraging by this species (Bellrose 1980). Blue-winged Teal distribution showed the same diagonal band component as that of Mallard with which it shares the ability to exploit more open habi- tats (Bellrose 1980). Numbers were too low, howey- er, to determine any relationship with either ecodis- trict or substrate type. Common Mergansers showed concentrations in those parts of the northern part of the Clay Belt that TABLE 2. Comparison of summary measures of waterfowl community parameters and habitat by ecodistrict and substrate. Classification Ecodistrict Substrate Significance Clay Moraine _ Significance (a) Waterfowl Community Parameters 29(n=81) 30 (n= 20) (p) m= 72) (n = 43) (p) Total Breeding Density 95.08 151.14 NS 115.00 97.46 NS (IP/100 km?) (0.094) Mean Number of Species per Plot 3.85 5.60 0.025 4.36 4.21 NS (b) Habitat Mean Area per Plot of All Wetland Habitat (Ha) 20.75 2797 0.003 21153 2V51 NS Number of Waterbodies per Plot | 4.45 0.012 3.68 3.21 NS 2002 Ross, ABRAHAM, GADAWSKI, REMPEL, GABOR, AND MAHER: BREEDING WATERFOWL 47 Mallard IP/100 km” CL] 0-5 [J 5-10 BB 10-20 HE 20-40 Mi 40-80 American Black Duck ore Green-winged Teal 1P/100 km O o-5 f] 5-10 HB 10-20 HB 20-40 Mi 40-20 Blue-winged Teal 1P/100 km” 2 * 1P/100 km American Wigeon IP/400 km? 0-5 5-10 10-20 20-40 Ring-necked Duck 4 Common Goldeneye IP/100 km DO o-s f 5-10 Hi 10-20 Hi 20-40 «40-80 Hooded Merganser eee Common Merganser oO 0-5 f 5-10 fH 10-20 & Hi 20-40 Hi «40-80 Number of Waterfowl Species per Plot Total Number of Waterfowl oe FiGURE 2. Contoured maps showing breeding densities of individual waterfowl species, overall breeding density, and species richness 48 THE CANADIAN FIELD-NATURALIST Vol. 116 - TABLE 3. Comparisons between ecodistricts and between substrates of breeding densities of the major waterfowl species in the Clay Belt. Classification Ecodistrict Division No. 29 No. 30 Species Mallard 25.51 20.00 Ring-necked Duck 12:35 oY 92 American Black Duck 13.81 29.47 Common Goldeneye 8.85 2125 Green-winged Teal 8.54 7.50 Hooded Merganser 6.69 6.67 Blue-winged Teal 5.45 6.25 Common Merganser 2.88 6.67 Canada Goose 2.06 8.33 American Wigeon ' 3.70 4.25 had high concentrations of Black Ducks and Ring- necked Ducks. No relationship with either substrate or ecodistrict could be demonstrated, probably due to the small numbers recorded. Common Mergansers have shown an affinity for rivers (McNicol et al. 1987; Rempel et al. 1997) in the Boreal Forest and their distribution in the Clay Belt may reflect the concentration of tributaries that converge toward the Moose River in the northeast. Small numbers of Canada Geese were found north of Lake Abitibi and Highway 11 in habitat showing strong muskeg elements of the Hudson Bay Low- lands which lie immediately to the north, and are the centre of the breeding range of the Southern James Bay Population of Canada Geese (Raveling and Lumsden 1977). Breeding density was significantly higher in Ecodistrict 30. American Wigeon were uncommon and not recorded in sufficient numbers to define distribution or determine regional differences. The species prefers larger permanent marshes with much open water (Palmer 1976) and shows an affinity for lake estuary marsh in the Clay Belt (Rempel et al. 1997). Northern river deltas held the highest breeding den- sities of this species (Bellrose 1980). Overall Waterfowl Distribution and Species Richness Contoured maps of total breeding pair density of waterfowl and numbers of waterfowl species per plot (Figure 2) indicated that both overall waterfowl den- sity and species richness were higher in the north. However, there was a significant relationship only between ecodistrict and species numbers (higher in Ecodistrict 30; Table 2). This may be an “edge effect” (i.e., higher habitat diversity) due to the inter- gradation with the Hudson Bay Lowlands immedi- ately to the north. Another area of higher species richness and density occurred around Timmins and Lake Abitibi near the south-eastern edge of the Clay Substrate Significance Significance (p) Clay Moraine (p) NS 29.17 14.92 0.013 < 0.001 20.95 17.05 NS 0.008 16.39 18.00 © NS < 0.001 12.38 10.85 NS NS 8.45 9.30 NS NS 5.09 9.88 0.04 NS 6.48 BWA!) NS NS 3.82 5.04 NS 0.006 2.89 3.29 NS NS 3.70 1.55 NS Belt, possibly reflecting greater habitat diversity where the Clay Belt grades into the Great Lakes-St. Lawrence Forest Region. Lowest species richness and density occurred around the southwestern edge in a very sandy section of the Chapleau Plain. Comparison with Other Studies Our survey showed that waterfowl breeding in the Clay Belt were not distributed uniformly. Species richness was significantly higher towards the north, particularly in Ecodistrict 30, and total numbers of breeding pairs showed a similar tendency. This gra- dient towards the north was significant for four species (Canada Goose, Black Duck, Ring-necked Duck, and Common Goldeneye), all of which are characteristic of boreal habitats and seem to respond to physiography, particularly the amount of wetland habitat. Mallards and Blue-winged Teals were con- centrated in a northwest-southeast strip bisecting the Clay Belt which correlates with the distribution of clay substrate. These species are more broadly dis- tributed in other habitats to the south and west and may be responding either directly or indirectly to the higher productivity associated with clay (Merendino and Ankney 1994). Other species, such as the American Wigeon and the Green-winged Teal, did not exhibit either of these patterns and instead may be responding to distributions of very specific habi- tat types. Initial comparison of our results with those of ear- lier surveys (Table 1) indicated substantial differ- ences. However, the previous surveys only covered part of the Clay Belt and locations of survey plots in earlier studies strongly influence the results. The Clay Belt plots used by Dennis (1974) were mostly in the relatively wetland-rich Ecodistrict 30 and other areas north of Highway 11, where one would expect a higher total breeding density than the aver- age from the present survey. Conversely, the survey by Ross (1987), that yielded a relatively low total 2002 Ross, ABRAHAM, GADAWSKI, REMPEL, GABOR, AND MAHER: BREEDING WATERFOWL 49 breeding density, was restricted to a 100 x 100 km block in the south central portion of the Clay Belt and included areas with the lowest breeding densities in this study. These relationships hold for three of the most common species. Black Duck and Common Goldeneye densities were highest in the Dennis (1974) survey while Black Duck density was lowest in the Ross (1987) count. Both species demonstrate the previously noted north-south density gradient. Densities for the Mallard, which did not show the north-south gradient, differed little among surveys. This implies that there have not been major changes in the Clay Belt populations of these three species since 1973. Of the more abundant species, only the Ring-necked Duck showed results [lowest count in the Dennis (1974) survey] that are not explainable by their present distribution. Total waterfowl breeding density is slightly higher on the Clay Belt than on the surrounding boreal forest characterized by exposed Precambrian Shield although the ranges of values partially overlap [Clay Belt (Table 1): 91.8-126.1 IBP/100 km? vs. Exposed Shield (Ross 1987): 75.0—107.8 IBP/100 km72]. Species composition, however, differs considerably. Generally, dabblers such as the Mallard, and Blue- winged Teal are more common in the Clay Belt whereas the mergansers attain higher densities on the exposed shield; goose numbers are minimal in both areas. The higher density of wetlands in the exposed shield appears to compensate in part for the higher productivity of individual wetlands in the Clay Belt (Ross 1987). The Hudson Bay Lowland immediately to the north supports, lower densities of ducks (approx. 37 IBP/100 km2; Ross, unpublished informa- tion) but higher numbers of geese (S—13 IBP/100 km? for Canada Geese). Overall the contribution of the Clay Belt to northern Ontario duck numbers is essen- tially proportional to its area. Conservation Implications Better understanding of distribution of waterfowl and characteristics of their breeding habitat allows more focussed and effective conservation actions. Areas being considered for acquisition and protection can be priorized based on waterfowl breeding density and species richness. Numbers of waterfowl] potential- ly at risk due to large-scale developments, (e.g., forestry, hydro-electric dams, and mining) can be determined and remedial action plans prepared. Furthermore, some insight can be gained into the effects of specific management actions. For example, efforts to increase the amount of wetland habitat may well show colonization by species such as Black Duck, Ring-necked Duck, Common Goldeneye, and possibly Canada Goose. If this managed habitat also has a rich clay substrate, one can also expect an increase in Mallards (and possibly Blue-winged Teal); however, this may lead to competitive interaction between Black Ducks and Mallards (Merendino and Ankney 1994). Target species and substrate type should be carefully considered in determining site locations for habitat management activities in the Clay Belt. Acknowledgments Financial support was provided by Ducks Unlimited Canada, the Ontario Ministry of Natural Resources through the Northern Forest Development Group, Wildlife Habitat Canada, and the Canadian Wildlife Service. We are very grateful for the field assistance provided by M. Barr, P. Burke, B. Di Labio, D. Fillman, D. Kopenhaver, D. Morgan, S. and S. O’Donnell, M. Porter, W. Simkin, N. Wilson, and J. Woodcock. J. Boos, R. Clay, and D. Kroeker of Ducks Unlimited staff at various times all made important contributions. We thank R. Watt of the OMNR for his support of this study and also the OMNR field offices whose radio operators provided flight following for our aircraft. We greatly appreci- ate the work of B. Campbell in data processing and figure preparation. Helpful comments were received from A. J. Erskine and an anonymous reviewer. Literature Cited Addy, C. E., W. F. Crissey, H. R. Webster, and G. F. Boyer. 1952. Waterfowl breeding ground survey in east- ern Canada. U.S. Fish and Wildlife Service Scientific Report Number 21. 81 pages. Bellrose, F. C. 1980. Ducks, Geese and Swans of North America. 3rd edition. Stackpole Books, Harrisburg, Pennsylvania. 540 pages. Chamberlain, E. B., and C. F. Kaczynski. 1965. Prob- lem in aerial surveys of waterfowl in eastern Canada. 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The Forest Resources of Ontario. Department of Lands and Forests, Toronto. Siegel, S. 1956. Nonparametric Statistics for the Behavioral Sciences. McGraw-Hill Book Company, New York. 312 pages. | Wellein, E. G., and H. G. Lumsden. 1964. Northern forest and tundra. Pages 67-76 in Waterfowl Tomorrow. Edited by J. P. Linduska. U.S. Department of the Interior, Washington, D.C. Wicken, E. 1986. Terrestrial ecozones of Canada. Eco- logical Land Classification Series 19. Environment Canada, Ottawa. 26 pages. Wickware, G. M., and C. D. A. Rubec. 1989. Ecoregions of Ontario. Ecological Land Classification Series Number 26. Environment Canada, Ottawa. 37 pages. Received 12 October 2000 Accepted 3 May 2002 Resilience of Foothills Rough Fescue, Festuca campestris, Rangeland to Wildfire EDWARD W. Bork!, BARRY W. ADAMS2, and WALTER D. WILLMS? !Range Scientist, Department of Agricultural, Food, and Nutritional Science, University of Alberta, 410E, AgFor Centre, Edmonton, Alberta T6G 2P5 Canada, 2Range Management Specialist, Public Lands Division, Alberta Agriculture, Food, & Rural Development, Bag 3014, Agriculture Centre, Gaol Road, Lethbridge, Alberta T1J 4CJ Canada 3Range Ecologist, Lethbridge Research Centre, Agriculture and Agri-Food Canada, P.O. Box 3000, Lethbridge, Alberta T1J 4B1 Canada Bork, Edward W., Barry W. Adams, and Walter D. Willms. 2002. Resilience of Foothills Rough Fescue, Festuca campestris, rangeland to wildfire. Canadian Field Naturalist 116(1): 51-59. A three year monitoring program evaluated the effects of a December 1997 wildfire in southwest Alberta, on Foothills Rough Fescue grassland species composition, ground cover, herbage production, and forage quality. Changes in species abundance included a reduction in grass cover (p<0.10) after burning. Rough Fescue also increased seedhead production during the second year after fire (p =0.08). Relative to the unburned area, graminoid production declined (p< 0.05) by approximately 40% with burning, while forb production was unaffected. By the second growing season, live plant cover and herbage production had recovered on the burned area. The forage quality of individual Foothills Rough Fescue plants was greater on the burned area, with the greatest increase in crude protein in 1998 (p<0.10), and energy and total digestibility in 1999 (p< 0.05). Increased quality may be linked to the level of forage production, as well as a fire-induced delay in plant phenology. Although soil erosion appeared to be minimal, there was an increase in exposed soil and a corre- sponding decline in litter and mulch cover (p< 0.05). Greater nitrogen levels (p= 0.051) were found in creeks downstream of the burn area during 1998, indicating some nutrient losses may be attributed to the fire. Although the grasslands exam- ined displayed considerable resilience to this severe wildfire, favourable recovery was probably linked to the high precipi- tation during 1998, when summer rainfall was 48% above average. Key Words: Foothills Rough Fescue, Festuca campestris, forage quality, precipitation, production, resilience, Alberta Foothills Rough Fescue (Festuca campestris of fescue grassland to fire have been conducted, these Rydb.) grasslands are well-adapted to fire, having a __ typically involve Plains Rough Fescue (Festuca hallii historical fire return interval of 5-10 years (Wright (Vasey) Piper) grasslands in the Aspen Parkland of and Bailey 1982). During the last century, however, Alberta and Saskatchewan (Bailey and Anderson fragmentation of rangeland landscapes and fire sup- 1978; Anderson and Bailey 1980; Redmann et al. pression is thought to have enhanced the risk of 1993; Gerling et al. 1995). Furthermore, these studies severe fires by lengthening fire return intervals and examine areas burned under prescription rather than increasing fuel accumulation during the burn inter- _ wildfire. Antos et al. (1983) examined changes in cession. Antos et al. (1983) concluded that Foothills species composition following a small (49 ha), uncon- Rough Fescue grasslands were best adapted to mod- __ trolled wildfire in July within a Foothills Rough erate frequency fire (e.g., every 6 years). Fescue-Idaho Fescue (Festuca idahoensis Elmer) From January of 1997 through August of 2000, plant community in Montana. eight known wildfires were documented in the As a result of concerns associated with increasing- Fescue Prairie region of Alberta affecting nearly ly common wildfire events in Alberta and the lack of 33 500 ha, with at least five of these greater than 400 information on their ecological impacts, a research ha (Barry Adams, personal observation). This trend plan was formulated to assess Foothills Rough prompted concerns over the ability of these grass- Fescue grassland recovery following a large-scale lands to recover from wildfire, particularly those fol- wildfire that occurred in December 1997. Initial lowing the prolonged absence of fire. Native rough inspection of the burned area indicated that many fescue grasslands may be less tolerant of infrequent rough fescue plants had been heavily damaged fire due to the increased severity of burning associat- and/or killed, presenting a unique opportunity to ed with litter accumulation. Rough fescue in particu- examine the impact of a dormant season fire on the lar, is of value to the ranches in the region because of _ resilience of these grasslands. Specific objectives of its role in providing a practical and economical this project were to determine the short-term (2-3 source of fall and winter grazing (Willms et al. year) impacts of the fire on: (1) plant community 1993), as well as habitat for wildlife. composition and ground cover, (2) above-ground net Although several studies investigating the response primary production, and (3) rough fescue forage 51 52 THE CANADIAN FIELD-NATURALIST quality. A secondary objective was to investigate the area for soil erosion. Methods Study Area The study area was burned by a wildfire on 14 December 1997, covering 220 km? southwest of Granum, Alberta (113°50”W; 49°45”N) within the Fescue Prairie Ecoregion (Strong 1992). The fire began around 11:00 and traversed 33 km in under four hours. The fire was preceded by a dry autumn and aided by sustained winds of 30—40 km/hr, gust- ing to 70 km/hr. Weather conditions at the time of the fire were unusual for December, with a maxi- mum mid-day temperature of 13°C (10°C above average) and relative humidity of 17% (Tymstra 1998*). Although detailed fuel-load data are unavailable for the burned area itself, litter loads adjacent to the burn averaged nearly 900 kg/ha. Approximately 83% of the 21 600 ha burn affected Foothills Rough Fescue grasslands. The economic and social impacts of this fire on the ranching com- munity within the region were widely publicized at the time. A fire intensity assessment prepared by Tymstra (1998*) described the Granum fire as extremely hot with a head fire intensity ranging from 10 000 to 20 000 kW/m2. Tymstra (1998*) further concluded that the average rate of fire spread (~10 km/hr) was one of the greatest documented for grassland fires in Canada. Topography of the area varied from steep foothills to gently sloping terraces, with occasional flat valley bottoms. Elevations range between 1 000 and 1 500 m. Native grasslands are dominated by mid to late successional Foothills Rough Fescue-Parry Oatgrass (Danthonia parryi Scribn.) communities, interspersed with riparian vegetation along wetlands and streams, and xeric grasslands dominated by Idaho fescue and Parry Oatgrass on sites with shallow or poorly devel- oped soils. In some areas, livestock grazing has resulted in an increase in introduced species such as Kentucky Bluegrass (Poa pratensis L.) (Willms et al. 1995, 1996). The dominant soil type is an Orthic Black Chernozem (Johnston et al. 1971). Field Procedures In May 1998, 10 sites were selected within Foothills Rough Fescue-Parry Oatgrass (Festuca- Danthonia) communities for monitoring, with eight situated along the perimeter burn as paired burned- unburned sampling sites, increasing the likelihood of evaluating vegetation responses to the fire itself. All paired sites were located on a uniform ecosite (i.e., aspect and soils) and were intersected by a human- made (i.e., grader-bladed) fire boundary. This strati- fication helped ensure that vegetational differences between paired transects were caused by the fire Vol. 116 ~ rather than a naturally occurring difference in physi- cal site characteristics. Two additional sites were sampled within the interior of the burn. At each site, a 30-m linear transect was randomly established and permanently marked to facilitate re- sampling in later years. On each transect, ocular esti- mates of canopy cover were done on all plant species within 15 systematically placed 0.1-m? quadrats (Daubenmire 1959) at peak vegetative growth in June of each year. In addition, the cover of loose litter and mulch, as well as the amount of bare soil were esti- mated. Mulch was defined as the matted layer of fine organic material overlying mineral soil, consisting of heavily degraded litter in combination with fine roots from plants, either dead or alive (Dix 1960). Herbage production was determined within four, 1.5 m by 1.5 m portable cages randomly set up along each transect. During the last week of August in each year, all current above-ground net primary produc- tion (ANPP) was clipped within a 0.5-m? quadrat in each cage. Cages were moved to new locations between years. All ANPP was sorted to graminoid, forb, and litter components, dried at 50°C for 3 days, and weighed to determine dry matter. Rough fescue seedhead counts were done the first week of August within 15, 1-m? quadrats nested over the Dauben- mire quadrats along each transect to assess the potential for seed production in each year. To evaluate the effect of burning on forage quali- ty, individual rough fescue plants were randomly selected and harvested during the first week of August. In 1998 and 1999, five and eight plants, respectively, were clipped on each transect. After standing dead material was removed, plant samples were analysed for crude protein (CP), acid detergent fiber (ADF), and total digestible nutrients (TDN), as these variables are important for evaluating the qual- ity of forage available to livestock and wildlife (Holechek et al. 1998). In the third growing season after burning, vegetation sampling was limited to seedhead counts, the evaluation of exposed bare soil, determination of ANPP, and litter mass. Watershed-level impacts of the fire were assessed from water samples taken within three creeks that flowed through the burn area. Samples were taken immediately above and below the burn between 22 January and 23 March, 1998 during snowmelt. All samples were analysed for total suspended solids (TSS), total dissolved sediment (TDS), and nitrogen (N) (Eaton et al. 1995). Data Analysis Direct comparisons were made using paired t-tests between the burned and unburned transects along the fire boundary on species richness (number of species), the cover of major grasses including foothills rough fescue and the wheatgrasses (Agropyron spp.), bare soil, litter, and mulch. The wheatgrasses [Northern 2002 (Agropyron dasystachyum (Hook.) Scribn.), Western (Agropyron smithii Rydb.), and Bearded (Agropyron subsecundum (Link) A.S. Hitchc.)] were pooled dur- ing sampling due to difficulty in identifying each species based on vegetative characteristics, particular- ly within the recovering burn area. Total legume cover was also assessed because these species constitute a key functional group responsible for N-fixation. Separate t-tests were done on the 1998 and 1999 data to evaluate the extent of recovery through time. Although data from the two transects in the interior of the burn could not be directly (i.e., statistically) com- pared with those from the burn perimeter, data from these sites were averaged for comparison to the other locations. Graminoid and forb ANPP, litter, and Rough Fescue seedhead counts, along with forage quality parameters, were also tested for burning effects in 1998, 1999, and 2000. Results and Discussion Growing Conditions Precipitation the year after the fire was favourable for recovery, with the majority falling during the growing season (Figure 1). Precipitation in 1998 at the nearby Agriculture Canada research station near Stavely was 502 mm from May to August, with a total of 647.5 mm in 1998, 46% above the regional average reported by Environment Canada (unpub- lished data). These data corroborate records from a ranch situated within the burn area, where April through August rainfall totalled 538 mm, and annual precipitation totalled 771 mm. In 1999, growing con- Bork, ADAMS, WILLMS: RESILIENCE OF FOOTHILLS ROUGH FESCUE 53 ditions approximated the norm for the region, with a total of 285 mm from May to August and annual pre- cipitation of 399 mm (Figure 1). Landscape patterns of green-up were variable in 1998, with some areas developing rapidly, as evi- denced by advanced vegetative growth and the emer- gence of Rough Fescue seedheads by early May. Plant development appeared to be particularly advanced at the perimeter of the burn, possibly due to increased soil temperatures associated with the removal of litter (Antos et al. 1983), which could hasten tiller development. In other areas, however, phenology was noticeably delayed. Variation in veg- etation development is likely attributed to differ- ences in ecosite conditions such as slope, aspect, and soils, as well as corresponding variation in fire behaviour across the landscape. General reconnai- sance suggested that the slowest development occurred within the centre of the burn where the greatest evidence of fire intensity was observed both during the fire (i.e., in terms of flame length and rate of spread) and after the burn (1.e., surface distur- bance). At these locations, intense heat may have penetrated more deeply into the soil, increasing dam- age to plant meristems and carbohydrate reserves by destroying plant tissue. Greater damage, in turn, would force plants to resume growth from perennat- ing buds, slowing growth. In 1999, the phenological development of burned vegetation was uniform throughout the burned area and more similar to that of the adjacent unburned vegetation. | i Jan.-April @ May-June OJuly-Aug. 0 Sept.-Dec. | c A ope o = 2 oO 2 a 1998 1999 2000 40 Year Ave FiGurE 1. Yearly precipitation from January 1997 to December 2000 for the Agriculture Canada Stavely sub-station, locat- ed approximately 50 km north of the December 1997 wildfire, and comparison to long-term average precipitation from the Claresholm—Meadow Creek, Alberta climate station (Environment Canada, unpublished data). 54 THE CANADIAN FIELD-NATURALIST Plant Species Abundance and Ground Cover A summary of the differences in plant species abundance in 1998 and 1999 among each of the three sample locations (unburned, perimeter burn, and interior burn) is provided in Table 1. Differences between burned and unburned sites along the fire boundary are consistent with the notion that fire is effective in changing plant community composition (Daubenmire 1968). Species richness (number/1.5 m2) and overall diversity were similar between burned and unburned areas (Table 1). Although the cover of introduced species tended to increase by the end of the second growing season (Table 1: 1999 data), this change was not significant (p> 0.10). Concerns that Kentucky Bluegrass, due to its strongly creeping growth habit, might be more tolerant of fire and capable of invad- ing areas where other plants had been removed by fire, did not appear to be substantiated. At two sites, Kentucky Bluegrass, and to a lesser extent Dandelion (Taraxacum officinale Weber), did increase indicat- ing the expansion of invasive species may be linked to their presence and abundance within the communi- ty prior to burning. Only total grass cover was reduced within the burned area relative to the unburned in 1998 (p<0.10; Table 1), with recovery by 1999. In com- parison to the perimeter burn, grass cover was less but forb cover greater, within the burn interior. Species data indicate the forb response was likely due to an increase in legumes such as Lupine (Lupinus argenteus Pursh), Locoweed (Oxytropis sericea Nutt.), Buffalo Bean (Thermopsis rhombifo- lia R.Br.), and Vetchling (Vicia americana Mubhl), all of which were more abundant in both sampling years (Table 1). Anderson and Bailey (1980) also found increases in 4 of 5 legume species associated with annual prescribed burning in Plains Rough Fescue grasslands of the Parkland in central Alberta. Other research has indicated legumes are well adapt- ed to fire and increase following burning due to the termination of seed dormancy (Martin et al. 1975). Although not tested here directly, this factor may contribute to the observations made in this study and merit further investigation within Foothills Rough Fescue grasslands. Regardless of the mechanism, the increase in legumes is important in that they con- tribute nitrogen through symbiotic N-fixation, which may in turn, facilitate plant community recovery fol- lowing fire. Of the dominant grasses, Parry oatgrass appeared to decline the most the year after fire, followed by the wheatgrasses (Table 1). In contrast, sedges (Carex spp.) and Foothills Rough Fescue appeared to be resilient, with minimal difference in canopy cover across the fireline (Table 1). Rough fescue, however, had notably less cover at the interior of the burn rela- tive to the perimeter. Assuming these transects had Vol. 116 similar amounts of Rough Fescue as the other areas sampled (this assumption is supported by the pres- ence of numerous dead fescue plants in the interior), burning may have been more severe at this location. The interior burn transects were generally located in areas where previous grazing may have been less intense, leading to greater litter accumulation. This interpretation is supported by the results of a 1990 range survey that described the grazing history of the interior burn area as very light, and was mapped as secondary range with considerable litter build up (Tannas 1990). With the exception of Parry Oatgrass, all domi- nant grasses appeared to recover by the second year. The resilience of Foothills Rough Fescue following burning is inconsistent with other studies document- ing fire-induced declines in rough fescue (e.g., Bailey and Anderson 1978; Antos et al. 1983). Most previous studies, however, have examined Plains Rough Fescue rather than Foothills Rough Fescue, which differ in morphology (Pavlick and Looman 1984), and presumably, their tolerance to fire. Mitchell (1957) suggested that the coarse stubble of rough fescue normally insulated perennating buds near the soil surface. With dry conditions, heavy stubble may allow the development of hot fires that burn into plant crowns as observed at the interior study locations. Antos et al. (1983) reported severe damage and high mortality of rough fescue on ungrazed sites where low fire frequencies allowed heavy litter accumulation, with three years needed for fescue recovery. In this investigation, the wheatgrasses declined more than Rough Fescue despite being generally considered better adapted to tolerate fire due to their rhizomatous growth habit (Wright and Bailey 1982). Other studies of wheatgrass response to burning show varied results, although most research has been done in drier regions. In some, fall burning increases wheatgrasses (Wright 1974; White and Currie 1983), while in others, wheatgrasses decline (Erichsen- Arychuk et al. 2002). The response observed here could also be due to the presence of Bearded Wheatgrass within the stand, which lacks rhizomes and may be susceptible to fire. Exposed Soil, Litter Cover, and Erosion Losses. Cover of bare soil, litter, and mulch varied with sampling location (Table 1). In general, litter was significantly (p<0.05) less across the fire-line on the burned area from 1998 through 2000, with a corre- sponding increase in bare soil and mulch. By 2000, bare soil was no longer different between adjacent burned (3.4%) and unburned areas (1.4%), likely due to plant recovery and the re-accumulation of litter. Bare soil was significantly (p <0.05) less across the fire boundary in 1999 but not 1998 (p>0.10), and may indicate some drying and loss of protective mulch on the burned area between sampling periods. 2002 Bork, ADAMS, WILLMS: RESILIENCE OF FOOTHILLS ROUGH FESCUE 55 TABLE 1. Mean canopy cover (%) of major plant species and functional groups at each location sampled in 1998 and 1999. Species listed only include those with a minimum average cover of 1%. Values in parentheses represent one SE for vari- ables with significance tests. Canopy Cover — 1998 Canopy Cover — 1999 Interior Burn Perimeter Burn Unburned Interior Burn Perimeter Burn Unburned Species Agropyron spp. 24 6.5 BA 12.4 15.5 11.9 Carex spp. £3 7 7.8 4.2 16.7 i343 Danthonia parryi 0.4 0.8 2 223 4.1 ke Bo Festuca campestris 3.6 10.1 11.6 25.5 22.4 23.5 Festuca idahoensis 4.1 5 0.2 13.9 53 Helictotrichon hookeri 0.9 0.8 5.2, at Koeleria macrantha 0.8 2.3 33 1.8 Poa pratensis 0.2 0.3 0:5 5.3 i 0.6 Poa sandbergii 4.2 2.4 Stipa viridula 0.2 2.3 4.8 Total Grass: 7.8 30.5a(3.8)* 47.5 b (5.6) 50.4 93.4a(4.8) 82.0a(7.8) FORBS: Achillea millefolium EZ Ta 0.1 Agoseris glauca 0.9 2.1 4.4 4.2 3.9 Androsace septentrionalis 6.2 0.6 Anemone multifida 1.1 2.8 7 zo | 4 (ws Aster laevis 0.6 0.3 0.1 | 0.1 0.6 Astragalus pectinatus 0.1 0.5 Hd | 0.9 1.6 Cerastium arvense 0.1 0.2 0.8 0.5 1.4 1.2 Commandra pallida 0.3 0.7 1.5 1.6 Companula rotundifolia 22 0.5 0.5 Erigeron caespitosum 22 ie? 6.3 2.1 Galium boreale ei 2.6 1.6 5 4.2 4.7 Geum triflorum 1.6 0.1 Ded, 0.3 Heterotheca villosa 1 3 1 3.4 2.8 Liatris punctata 0.3 0.8 i! 1.8 Lomatium dissecta 0.2 0.3 0.3 0.7 0.3 1.1 Lupinus argenteus 8.8 13 1.8 16.2 at 4.3 Oxytropis sericea 0.8 0.7 Ee 1.4 Phlox hoodii 0.1 0.7 0.4 lh Solidago missouriensis 0.1 0.7 2.8 0.9 Taraxacum officinale 0.1 0.3 0.4 0.4 0.7 0.3 Thermopsis rhombifolia 10.4 2.8 4.5 21.4 10.7 9.3 Tragopogon dubius 0.1 0.2 0.1 0.2 0.1 . Vicia americana 0.6 es 0.4 0.1 2 1.3 Viola canadensis 0.7 0.1 0.4 ] Zygadenus venenosus 0.1 0.9 0.2 0.6 1.3 0.1 Total Forb: 28.7 20.6a(4.5) 22.2 a (3.6) 82.1 55.9a(5.4) 46.8a(5.6) Legume Cover: 19.8 6.7 a (1.6) 8.5 a (1.8) 42.7 19.7a(4.3) 18.5a(3.9) SHRUBS: Artemisia frigida 0.3 2 1.4 2.3 Rosa arkansana eS 0.5 0.2 3.9 l 0.5 Total Shrub: 2,7 0.8 a (1.2) 2.2 a (1.3) 3.9 2.4 a (0.6) 3.1 a (0.4) INTRODUCED COVER 0.2 0.6 a (0.5) 0.9 a (0.4) 3.6 4.0 a (2.4) 1.0 a (0.5) RICHNESS (no./1.5 m2) 17.5 23.28a(1.8) 24.3 a(1.3) 21 27.8a(1.4) 26.0 a(1.5) SHANNON DIVERSITY 0.95 1.08 a (0.04) 1.08 a (0.04) 0.99 1.16a(0.04) 1.14 a (0.04) OTHER: Litter Cover 11.2 39.6 A (10.2) 91.7 B (5.8) 55.4 76.3 A (4.1) 97.9 B (1.7) Mulch Cover 76 52.1 A (8.0) 7.5 B (5.6) 28.2 14.7A (3.3) 0.0B(0) Bare Soil 11.9 5.4 a (3.6) 0.1 a (0.1) 15 6.6A(1.4) 0.3 B (0.2) Exposed Rock 1.3 0.8 a (0.4) 0.2 a (0.1) 1.2 0.5a(0.3) 0.1a(0.03) Microphytic Cover 2.9 a (2.1) 0.5 a (0.3) 1.9a(1.9) 1.8 a(1.7) *Within a component and year, cover values across the fire boundary with different upper and lowercase letters are signifi- cantly different from one another at p < 0.05 and p< 0.10, respectively. 56 THE CANADIAN FIELD-NATURALIST Overall, the greatest amounts of bare soil and mulch were evident at the interior burn, further supporting the notion that this area was more severely burned. Potential for soil erosion was a serious concern to the local community and land administration agen- cies. Soil drifting was so severe on adjacent cropland immediately following the burn that emergency tillage was imposed. In contrast, there was little or no evidence of mineral soil loss on burned range- land. Soil exposure was less than 1% on unburned areas, with 5.5 to 6.6% exposure on the perimeter burn and 11.9 to 15% for the interior burn during 1998 and 1999, respectively. In Foothills Rough Fescue grasslands, soil erosion by water tends to increase when soil exposure exceeds 15% (Johnson 1962; Naeth et al 1991); although soil exposure on the burned area increased, it failed to exceed this threshold. Comparative assessment of water quality above- and down-stream of the fire indicated that although total suspended and dissolved solids differed little, greater nitrogen was present in water collected from creeks immediately below the burned watersheds (Table 2; p=0.051). Thus, fire likely contributed to nitrogen loss during snowmelt and/or spring rainfall from the watersheds. Herbage ANPP and Litter Graminoid and forb ANPP followed patterns simi- lar to that for plant cover (Table 3). In particular, graminoid production declined (p< 0.05) by about 40% across the fire boundary in 1998, but remained negligibly less in 1999, with full recovery in 2000. In contrast, forb production appeared to be greater on the burned area in the years after the fire (Table 3), although this difference was not statistically sig- nificant (p>0.10). The greatest visible effect of burning in each year was at the interior burn, where a further decline in graminoid production and increase in forb production initially occured relative to the perimeter burn. Interestingly, this trend reversed in 2000, with graminoid and forb produc- tion apparently greater at the interior burn. The initial decrease in graminoid ANPP after fire is consistent with other studies for both Foothills Rough Fescue (Jourdonnais and Bedunah 1990) and Plains Rough Fescue grasslands (Redmann et al. 1993; Gerling et al. 1995). Recovery of production Vol. 116 in all these studies took at least two years. Redmann (1978) attributed declines in production following burning to increased plant water stress. However, the favourable precipitation during 1998 in the current study suggests the reduction may be from plant responses directly caused by fire rather than a change in soil water regime. Coupland (1974, as cited in Redmann 1978) also documented reduced productivity after fire in Mixed Prairie despite above-normal precipitation. Regardless of the mech- anism, the magnitude of production decline observed here may be linked to the dormant season timing of the burn, as autumn fires are more detrimental than spring fires (Redmann et al. 1993). Several other factors may contribute to the reduc- tion in graminoid production. Although the plant communities examined were in good to excellent range condition at the time of the fire based on the composition of unburned areas, drought or heavy grazing prior to the fire may have increased stress on graminoids and reduced their vigor or winter hardi- ness. Additive negative effects on vegetation have been shown between defoliation and drought (e.g., Hendrickson and Berdahl 2002) and defoliation and fire (e.g., Bunting et al. 1998). Although the com- bined impacts of defoliation and fire have been investigated on Foothills Rough Fescue (Bogen 2001), that study examined defoliation after burning rather than before. The unusually mild autumn preceding the Granum fire may also have allowed vegetation to continue development into December. Erichsen-Arychuk et al. (2002) documented variable grassland recovery in Dry Mixed Prairie landscapes following August wildfire under drought conditions, with landscape- based differences potentially due to the stage of plant development at the time of fire. Following burning, the reduction of insulation through the loss of litter and associated snow cover would result in colder soil temperatures in winter (Johnston et al. 1971), increasing the susceptibility of burned plants to freezing (Kowalenko and Romo 1998). As expected, litter mass was markedly less (p < 0.05) on the burned area (Table 3). Although the favourable growth during 1998 increased litter on the burn by the end of 1999, it remained less com- pared to the unburned area, with this trend continu- TABLE 2. Results of the analysis of water sampled from three creeks above and below the fire affected watersheds in 1998 and analyzed for total suspended solids (TSS), total dissolved sediments (TDS), and nitrogen (N). Mean per Sampling Location (SE): Significance*: Variable: Above Burn (n = 11) Below Burn (n = 11) T- statistic Probability TSS (ppm) 178.5 (44.5) 297 (152.4) 0.73 p = 0.482 TDS (ppm) 564 (56.4) 1010 (328.7) 1.42 p = 0.186 N (ppm) 115 (0.15) 3.88 (1.30) peer) p=0.051 *Paired t-tests contrast samples collected from above and below the burned portions of the watershed. 2002 Bork, ADAMS, WILLMS: RESILIENCE OF FOOTHILLS ROUGH FESCUE a7 TABLE 3. Mean (SE) current annual graminoid and forb production, as well as litter levels (kg/ha) within the unburned, perimeter burn, and interior burn locations in 1998, 1999, and 2000. Component Year Unburned Perimeter Burn Interior Burn Graminoid 1998 1466 (126) A* 940 (77) B 714 1999 1596 (154) 1498 (226) 1220 2000 966 (109) 844 (48) 1349 Forb 1998 439 (44) 545 (73) 682 1999 453 (37) 621 (105) 921 2000 110 (15) 133129) 292 Litter 1998 898 (69) 0 (0) 0 1999 3351 (761) A 852 (98) B 662 2000 2101 (381) A 908 (28) B 948 *Within a component and year, comparisons between burned and unburned means across the fire boundary represented by different letters are significantly different (p < 0.05). ing into 2000 (Table 3). Litter is important for con- serving soil water and maintaining herbage produc- tion (Weaver and Rowland 1952; Willms et al. 1986). Antos et al. (1983) found the loss of litter after fire increased soil temperatures, reducing the near-surface effective water regime. Fluctuations in litter, even on the unburned area (Table 3), indicate how rapidly this variable can change. Litter is lost as a result of fire, herbivory, microbial decomposition, and weathering, and is affected by grassland species composition (Facelli and Pickett 1991). Willms et al. (1996) found 23% of total biomass disappeared between fall and spring on good condition rough fes- cue grassland, while 56% disappeared from stands dominated by forbs and introduced grasses. Given the slow re-accumulation of litter in this study, it appears several years are needed for litter re-accu- mulation following the Granum fire. Rough Fescue Seedhead Production Burning increased (p< 0.10) Rough Fescue seed- heads two years later, when seedhead densities on the burned area were more than twice that of the unburned area (Table 4). The delay in response fol- lowing disturbance is similar to that found in other studies for Foothills Rough Fescue (Johnston and MacDonald 1967; Willms 1988). Gerling et al. (1995) found Plains Rough Fescue increased seed- head production the year following fire, but only when burned in spring or early summer (i.e., 1 June or earlier) of the previous year rather than in late summer or fall. Burning may have increased seedhead production by the addition of nutrients to the soil or the loss of litter. Litter removal may trigger seedhead establish- ment through increases in photosynthetically active radiation (PAR) within the meristematic region of the plant crown (Willms 1988). The delayed response indicates that either plant stress immediate- ly after burning inhibited reproduction the first year, or more likely, that there is a time lag needed for developing tillers to become reproductive, as deter- mined by the new environmental (e.g., light and/or nutrient) conditions after fire. It should also be noted that seedhead production was generally greater on the unburned area in 1999 (19.5 + 3.8) than in 1998 (2.9+ 1.1) (Table 4). Thus, abundant seedhead pro- duction in 1999 on the burned area is due to the combined effect of favourable moisture during 1998, coupled with burning. The increase in seedheads reflects an important ecological adaptation of Foothills Rough Fescue to ensure this bunchgrass recolonizes areas of soil following burning, and thus, ensures its perpetuation within the plant community. Rough Fescue Forage Quality Burning positively affected the forage quality of Rough Fescue. In particular, crude protein and total digestibility increased, while acid detergent fiber (ADF) decreased (Table 5). However, between-plant variation was considerable, with only crude protein differing significantly (p < 0.10) across the fire boundary in 1998 (other variables had p< 0.15). TABLE 4. Rough Fescue seedhead production on the three sampling areas from 1998 to 2000. Seedhead Densities — #/m2 (SE): Year Interior Burn Perimeter Burn 1998 0.2 (0.1) 1.1 (0.6) 1999 56.2 (17.3) 42.0 (10.9) 2000 0.4 (0.1) 0.1 (0.1) Significance*: Unburned T- statistic Probability okt hol) 2.21 p=0.11 19.5 (3.8) 2.64 p=0.08 0.8 (0.4) Ete p=0.18 *Paired t-tests contrast data collected across the fire boundary (i.e. Perimeter Burn vs Unburned). 58 THE CANADIAN FIELD-NATURALIST Vol. 116 - TABLE 5. Mean (SE) percent crude protein (CP), acid detergent fiber (ADF), and total digestible nutrients (TDN) of Foothills Rough Fescue plants sampled within the unburned, perimeter burn, and interior burn locations in 1998 (n =5 per transect) and 1999 (n = 8). Component Year Unburned Perimeter Burn Interior Burn CP (%) 1998 7.20 30) Ga. * 8.40 (0.44) b 10.10 1999 7.39 (0.29) 6.54 (0.36) 8.49 ADF (%) 1998 41.8 (0.7) 40.2 (0.5) 38.6 1999 42.2(0.4) A 40.9 (0.4) B 38.9 TDN (%) 1998 5221 (OS) 54.5 (0.3) 56.9 1999 S1.4(03)" A 53.4 (0.3) B 56.4 *Within a nutrient, comparisons between burned and unburned areas across the fire boundary represented by different uppercase (p< 0.05) or lowercase letters (p < 0.10) differ significantly. When larger sample sizes of plants were used in 1999, energy and total digestibility were greater (p< 0.05), and ADF lower (p < 0.05), within the burned area. Crude protein was unaffected (p > 0.10) in 1999, although it remained particularly high in plants sampled from the interior burn. The initial increases in protein are similar to the changes reported for Plains Rough Fescue following fire (Redmann et al. 1993) and burned Bluebunch Wheatgrass (Agropyron spicatum (Pursh) Scribn.) (Willms et al. 1981). Improved quality may be due to nutrient release into the soil following burning (Daubenmire 1968) or reduced competition on the burned area corresponding with a decline in live plant cover (Table 1). Increased quality may also stem from a delay in plant phenology throughout the summer, which would result in greater comparative quality at a fixed date of sampling. Delayed phenolo- gy of rough fescue grassland following autumn burn- ing was documented by Redmann et al. (1993), as was an associated increase in N concentration. Conclusions The 1997 Granum wildfire in Fescue Prairie had variable effects on Foothills Rough Fescue grassland plant species abundance and ground cover, herbage production, and forage quality, as determined by dif- ferences between the unburned and perimeter burn locations. Grass cover and production declined the year after fire, but recovered by 1999, likely as a result of the above average precipitation in 1998. Drought after burning may have had an additive detrimental impact on vegetation recovery, particu- larly in the absence of litter to conserve soil water. Declines in production also coincided with increases in the forage quality of Rough Fescue. The observed responses highlight the impacts of dormant season wildfire on Foothills Rough Fescue grasslands, including the resilience of the dominant species. Additionally, it should be noted that although species richness and diversity were not increased by fire in this investigation, there was evidence that fire was effective in increasing the abundance of forbs, particularly legumes. Thus, the importance of occa- sional fire in maintaining biodiversity should not be discounted within these grasslands. For many ranches affected by the fire, the need to restore litter through conservative grazing may seem to conflict with their natural concern about grass litter as a fire hazard. Balancing the need for litter to con- serve water and protect the range resource, yet pre- vent excessive fuel loading, will be an ongoing ques- tion for land managers to address. Fortunately, the more enduring negative impacts of fire appear to be confined to a relatively small area. The results of this project should also be of interest to the managers of protected areas where grazing and the normal cycling of biomass by large herbivores and fire have been altered from their natural process on the landscape. Acknowledgments This research was funded by the Alberta Cattle Commission (ILO Grant# 98-0987), with additional support from the University of Alberta, AAFRD- Public Lands, AAFC-Lethbridge, and Norwest Labs. The assistance of Amanda Bogen, Darlene Moisey, Janna Wowk, and Dianne Nadeau with field setup and data collection is greatly appreciated. This research was made possible by the cooperation of many ranchers within the area affected by the Granum wildfire. Their interest and dedication is greatly appreciated. Documents Cited (marked * in text) Tymstra, C. 1998. The 1997 Granum fire in southwest Alberta: A case study. Unpublished Alberta Environ- mental Protection Report, Land and Forest Services, Forest Protection Branch. 10 pages. Literature Cited Anderson, H. G., and A. W. Bailey. 1980. Effects of annual burning on grassland in the aspen parkland of east-central Alberta. Canadian Journal of Botany 58: 985-996. . Antos, J. A., B. McCune, and C. Bara. 1983. The effects of fire on an ungrazed western Montana grassland. American Midland Naturalist, 110: 354-364. Bailey, A. W., and M. L. Anderson. 1978. Prescribed burning of a Festuca-Stipa grassland. Journal of Range Management 31: 446-449. 2002 Bogen, A. D. 2001. Festuca campestris Rydb. response to fire and defoliation. M.Sc. thesis, Department of Agri- cultural, Food, and Nutritional Science, University of Alberta. 109 pages. Bunting, S. C., R. Robberecht, and G. E. Defosse. 1998. Length and timing of grazing on postburn productivity of two bunchgrasses in an Idaho experimental range. International Journal of Wildland Fire, 8: 15—20. Daubenmire, R. 1959. A canopy-coverage method of vegetational analysis. Northwest Science 33: 437-443. Daubenmire, R. 1968. Ecology of fire in grasslands. Pages 209-267 in: Advances in Ecological Research. Edited by J. B. Cragg. Academic Press, New York and London. Dix, R. L. 1960. The effects of burning on the mulch structure and species composition of grasslands in west- ern North Dakota. Ecology 41: 49-56. Eaton, A. D., L. S. Clesceri, and A. E. Greenberg. Editors. 1995. Standard methods for the examination of water and wastewater. 19th edition. American Public Health Association, Washington, D.C. Erichsen-Arychuk, C., E. W. Bork, and A. W. Bailey. 2002. Northern dry mixed prairie responses to summer wildfire and drought. Journal of Range Management 55: 164-170. Facelli, J. M., and S. T. A. Pickett. 1991. Plant litter: its dynamics and effects on plant community structure. Botanical Review 57: 1-32. Gerling, H.S., A. W. Bailey, and W. D. Willms. 1995. The effects of burning on Festuca hallii in the parklands of central Alberta. Canadian Journal of Botany 73: 937-942. Hendrickson, J. R., and J. D. Berdahl. 2002. Inter- mediate wheatgrass and Russian wildrye responses to defoliation and moisture. Journal of Range Management, 55: 99-103. Holechek, J. L., R. D. Pieper, and C. H. Herbel. 1998. Pages 286-330 in: Range management: Principles and practices. 3rd edition. Prentice Hall, Upper Saddle River, New Jersey. Johnston, A. 1962. Effects of grazing intensity and cover on the water-intake rate of fescue grassland. Journal of Range Management 15: 79-87. Johnston, A., and M. D. MacDonald. 1967. Floral initia- tion and seed production in Festuca scabrella Torr. Canadian Journal Plant Science 47: 577-583. Johnston, A., J. F. Dormaar, and S. Smoliak. 1971. 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Grazing impacts on litter and soil organic matter in mixed prairie and fescue grassland ecosystems of Alberta. Journal of Range Management 44: 7-12. Pavlick, L. E., and J. Looman. 1982. Taxonomy and nomenclature of rough fescues, Festuca altaica, F. campestris (F. scabrella var. major), and F. hallii, in Canada and the adjacent part of United States. Canadian Journal of Botany, 62: 1739-1749. Redmann, R. E. 1978. Plant and soil water potentials fol- lowing fire in a northern mixed grassland. Journal of Range Management, 31: 443-445. Redmann, R. E., J. T. Romo, B. Pylypec, and E. A. Driver. 1993. Impacts of burning on primary produc- tion of Festuca and Stipa-Agropyron grasslands in cen- tral Saskatchewan. American Midland Naturalist 130: 262-273. Robberecht, R., and G. E. Defosse. 1995. The relative sensitivity of two bunchgrass species to fire. Inter- national Journal of Wildland Fire 5: 127-134. Strong, W. L. 1992. Ecoregion and ecodistricts of Alberta. Volume 1. Alberta Forestry, Lands, and Wildlife, Land Information Services, Edmonton. 77 pages. Tannas, C. 1990. Range survey of selected grazing leases in the Southern Porcupine Hills. Alberta Forestry, Lands and Wildlife. Public Lands Division Report. 125 pages. Weaver, J. E., and N. W. Rowland. 1952. Effects of nat- ural mulch on development, yield, and structure of native grassland. Botanical Gazette 114: 1-19. White, R. S., and P. O. Currie. 1983. Prescribed burning in the Northern Great Plains: Yield and cover responses of three forage species in the mixed grass prairie. Journal of Range Management 36: 179-183. Willms, W. D., A. W. Bailey, A. Mclean, and C. Kalnin. 1981. Effects of fall clipping or burning on the distribu- tion of chemical constituents in bluebunch wheatgrass in spring. Journal of Range Management, 34: 267-269. Willms, W. D., S. Smoliak, and J. F. Dormaar. 1985. Effects of stocking rate on a rough fescue grassland veg- etation. Journal of Range Management, 38: 220-225. 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Received 23 October 2000 Accepted 21 March 2002 Survival, Fates, and Success of Transplanted Beavers, Castor canadensis, in Wyoming Mark C. MCKINSTRY and STANLEY H. ANDERSON Wyoming Cooperative Fish and Wildlife Research Unit!, Box 3166, Laramie, Wyoming 82071 USA McKinstry, Mark C., and Stanley H. Anderson. 2002. Survival, fates, and success of transplanted Beaver, Castor canaden- sis, in Wyoming. Canadian Field-Naturalist 116(1): 60-68. Beaver (Castor canadensis) through their dam building activities, store water, trap sediment, subirrigate vegetation, and subsequently improve habitat for fish, wildlife, and livestock. Many landowners realize the benefits that Beaver can bring to a riparian area and are interested in using them to improve this habitat. From 1994 to 1999 we trapped and relocated 234 Beaver to 14 areas throughout Wyoming to improve riparian habitat and create natural wetlands. We attached radio trans- mitters to 114 Beaver and subsequently determined movements and mortality of released Beaver, and the overall success of our releases. Mortality and emigration (including transmitter failure) accounted for the loss of 30% and 51%, respectively, of telemetered Beaver within 6 months of release. Kaplan-Meier survival estimates were 0.49 (SE = 0.068) for 180 days and 0.433 (SE = 0.084) for 360 days, and did not differ significantly between age classes. On average, 17 Beaver were transplanted to each release site, and at 11 locations, in an attempt to augment single Beaver that had become established and increase transplant success, we transplanted Beaver in two or more years. Success of an individual Beaver’s relocation was unrelated to any of the variables we tested, ‘although 2—3.5 year-old Beaver had higher average success (measured in days of occupancy at the release site) than older animals. Animals < 2 years old had 100% mortality and emigration losses within 6 months of release. High predation and mortality rates of our released Beaver may be due to habitat (our streams were shallow with no ponds and provided little protection) and extensive predator communities. We established Beaver at 13/14 of our release sites and they eventually reproduced. Our results show that Beaver can be relocated successfully but losses from mortality and emigration need to be considered and planned for. Key Words: Beaver, Castor canadensis, transplanting, reintroduction, translocation, predation, Wyoming Beaver (Castor canadensis) alter riparian-stream ecosystems through their woodcutting and dam- building activities. Beaver dams create a lentic habi- tat in an otherwise lotic system. These ponds retain sediment and organic matter in the channel, create and maintain wetlands, modify nutrient cycling and decomposition dynamics, modify the structure and dynamics of the riparian zone, alter hydrologic regimes (Butler 1991), and influence the character of water and materials transported downstream (Naiman et al. 1988). The resultant habitats are rich mosaics of diversity that are beneficial hydrological- ly (Hanson and Cambell 1963; Rabe 1970; Johnson et al. 1992), biologically (Jenkins and Busher 1979; reviewed by Hill 1982; Olson and Hubert 1994; Brown et al. 1996; McKinstry and Anderson 1999; McKinstry et al., 2001), and socially (Naiman et al. 1988). It is estimated that Beaver have been extirpat- ed from over 25% of the streams in Wyoming, and in many streams where they are still present their numbers have been reduced to where they are eco- logically absent (McKinstry et al. 2001). The elimi- 'The Unit is jointly supported by the University of Wyoming, Wyoming Game and Fish Department, U.S. Geological Survey-Biological Resources Division, and the Wildlife Management Institute. 60 nation of Beaver from portions of its historic range has been cited as a major influence on the structure and patterns of vegetation in these systems (Neff 1957; Barnes and Dibble 1986; Naiman et al. 1986; Nummi 1989; Kay 1994; Nolet et al. 1994). Throughout the intermountain west, interest has been expressed in improving riparian areas for wildlife, livestock, and humans (Apple et al. 1985; McKinstry and Anderson 1999; McKinstry et al. 2001). Beaver, through their dam building activities, can increase water storage, reduce sedimentation, and improve vegetation communities (Naiman et al. 1988), all of which are valuable to many landowners. Livestock are also attracted to Beaver-influenced areas for water, shade, and vegetation that remains green after upland forage has dried out. Forage pro- duction near these wetlands is often two to three times higher than comparable upland ranges (Apple et al. 1985; Chaney et al. 1991: 31). Many states have undertaken Beaver transplant programs to improve riparian areas (Smith 1980; Hill 1987; Butler 1991; Collins 1993; Vore 1993; McKinstry and Anderson 1997; McKinstry 2001) and managers with the Wyoming Game and Fish Department (WG&FD) decided to investigate the feasibility of a Beaver relocation program in areas where beaver have not recolonized due to isolation from dispersing populations and poor habitat conditions. In 1994 we initiated research to (1) document the 2002 effects of Beaver on riparian areas in Wyoming (reported in McKinstry et al. 2000; 2001), (2) assess Beaver management concerns from both private and public landmanagers (reported in McKinstry and Anderson [1999]), and (3) evaluate a Beaver reintro- duction project for the purpose of wetland creation and riparian improvement. Our objectives in this paper are to examine survival, mortality, emigration, and success of Beaver translocated in Wyoming for the purpose of riparian restoration. Study Area Beaver were trapped at 33 various locations in Wyoming (described in McKinstry and Anderson 1998) and translocated to 14 different 1s'-3™ order streams (< 0.283 m/sec) throughout Wyoming (Figure 1, Table 1). All drainages were walked a minimum of 3 km in both directions from the pro- posed release site to document any past or current Beaver activity. At 13 of the release sites, old Beaver sign (20-100 yrs) was present but we found no fresh activity. At the remaining site, Breteche Creek, prior Beaver activity was not found. All release sites had sufficient vegetation to support Beaver. Four streams were ephemeral and dry in early August each year [Ff a x? . Ps ie e a ea MCKINSTRY AND ANDERSON: TRANSPLANTED BEAVERS IN WYOMING 61 that we checked them (1993-1999), the remainder were perennial and carried water year-round (Table il) Methods Beaver were trapped using snares and Hancock traps (McKinstry and Anderson 1998) from areas where they were causing damage to landowners (pri- marily irrigation conflicts) (30 sites) or where they were so plentiful that selective removal would not significantly impact the habitat (3 sites). All Beaver were trapped from colonies that were dam and lodge builders (creek Beaver) as opposed to bank-denning non-dam builders (river Beaver). We felt that these animals would be more likely to create the desired habitat. We began trapping after ice-off in the spring (usu- ally early to mid May) and concluded trapping by the second week in October, depending on snow and ice conditions. All traps were set between 1600 h and 1900 h each day and were checked by 1000h the following morning to minimize the time that ani- mals spent in traps. On average we set 17 traps/ night/trapper; more than that and it was difficult to get them checked by 1000 h and find them in the ue 4 be nas rd i “a Vs Z =] =) = — = i=) D oe She : AY od : en Ye “f ’ than SSE ee: a ee rer Tau, mse not _ Ficure |. Beaver relocation sites in Wyoming. Letters correspond to release site locations described in Tables | and 2. | - : . | : ; 62 THE CANADIAN FIELD-NATURALIST - Vol. 116 TABLE |. Latitude and longitude, dominant vegetation, and stream classifications (using Rosgen’s [1994] classification) for Beaver release locations in Wyoming. Letters for release sites correspond with locations in Figure 1. Stream Name (nearest town) Latitude, Longitude Dominant Vegetation Stream Classification (A) Bear Gulch (Story) 44°31'46"N, 106°53'11"W Aspen and cottonwood DAS (B) Breteche Creek (Cody) 44°24'36"N, 109°23'28”"W Aspen and cottonwood A4 (C) Bush Creek (Shell) 44°27'41"N, 107°37'30"W Cottonwood A4 (D) Currant Creek (Rock Springs) 41°12'55”N, 109°22'20”"W Willow G5 (E) Deep Creek (Sundance) 44°47'29"N, 104°20'41"W Gambel’s oak and aspen (F) Ennos Creek (Thermopolis) 43°54'50"N, 108°54'14"W Willow B4 (G) Lake Creek (Saratoga) 41°28'16"N, 106°38'20”"W Narrowleaf cottonwood (Populus angustifolia) and willow (Salix sp) B4 (H) Little Red Creek (Casper) 42°42'55’"N, 106°23'48”W Cottonwood F4 (I) Prairie Dog Creek (Big Horn) 44°35'36"N, 106°54’30”W Aspen and cottonwood AS5 (J) S. Pine Creek@ (Sundance) 44°44'29"N, 104°20'30”W Scrub oak and common chokecherry (Prunus virginiana) F6 (K) Red Creek (Rock Springs) 41°04'01”"N, 109°02'30”W Engelmann spruce (Picea engelmannii) and cottonwood F5 (L) Spring Creek (Centennial) 41°12'59"N, 106°07'44"W ~~ Willow F5 (M) Trabing Creek (Big Horn) 44°36’01”"N, 106°58'59"W Aspen and western hawthorn (Crataegus succulenta) A4 astreams were ephemeral and dry by early August each year we checked (1993-1999) thick willows where we were trapping. Beaver were sexed through cloacal examinations (Larson and Taber 1980:163), weighed to determine age class (kit, yearling, sub adult, and adult), and ear tagged in both ears with small monel ear tags for identifica- tion. Beaver were held up to five days post-capture in a 2.2 by 3.1 m cage that allowed them free access to water; this was necessary since we only caught a few (<3) animals each day and we wanted to move them as a group to the release site which was always > 160 km away. Forty-six Beaver (range 11-31 kg) were implanted with Advanced Telemetry Systems (ATS, Isanti, MN) model 17 internal telemetry transmitters using techniques described by Davis et al. (1984). Implanted Beaver were monitored for a minimum of 24 hours post surgery prior to release (Davis et al. 1984). Beginning in 1998, 67 Beaver (range 10-25 kg) were outfitted either with tail collars (8 Beaver) or transmitters mounted on the tail using modified ear-tag transmitters (59 Beaver) (Roth- meyer et al. 2002) in an attempt to reduce logistical problems associated with surgeries conducted in the field (e.g., cost, time, sterility concerns). All radio transmitters were equipped with 24-hour mortality sensors. Animals were released after 1500 hours in an attempt to decrease predation during daylight hours. Beaver were monitored for movements and mor- tality for 2 days after release and approximately every 2 to 4 weeks thereafter. Mortality dates were calculated as the mid-date between the date found and the date of last-live location. At all release sites, streams were walked (minimum of 5 km) and flown (minimum of 10km) in both directions 3 months after release to determine if Beaver were established. In four instances non-transmittered Beaver became established, and walking the streams allowed us to look for evidence of tree cuttings or dam construc- tion and thus determine if non-telemetered Beaver were active. Since we were interested in using Beaver to improve habitat within 3 km of the release sites (usually headwater areas), we defined emigra- tion as Beaver that moved further than 3 km from the release site. Beaver moving > 5 km from the release sites were not monitored unless they built dams and lodges and remained stationary. Cause of mortality was determined through physi- cal examination of hair and scat samples found at the kill site (Moore et al. 1974), bite marks and subcuta- — neous bleeding (indicates animal was alive when bit- ten) on Beaver carcasses, track marks (O’Gara 1978), and, beginning in 1998, DNA analysis of hair and scat samples. Dr. Elizabeth Williams, Pathologist at the Wyoming State Veterinary Lab, performed lab work and necropsies. Tom Moore and Deedra Hawk, Forensic Supervisor and Research Associate, respectively, Wyoming Game and Fish Department, examined hair and scat samples. We used a z test (Jandel Scientific 1994) to test proportional differences in captures of males versus females. For estimates of survival we used Kaplan- Meier product limit estimators (as reviewed by White and Garrott 1990) for both 360 and 180 day survival rates. We used the Kaplan-Meier approach since our data was staggered entry and we had con- 2002 MCcKINSTRY AND ANDERSON: TRANSPLANTED BEAVERS IN WYOMING 63 siderable loss of individuals due to emigration or transmitter failure. To model success of transplants we used both logistic and multiple regression (White and Garrott 1990). For our logistic regres- sion models we coded each Beaver as either a O for failure (emigration or mortality) or a 1 for success (lived > 6 months, constructed a dam and lodge, and had the opportunity to reproduce). For multiple regression models we used the length of success (remained within 3 km of release site) in days as our dependent variable. We used weight, sex, age class, season of release (spring: May 9—June 15) or fall August 15-October 10), year of release (1994-1999), and number of cohorts released con- currently with each Beaver as our covariates. For age class we plotted weights and found natural dis- tinctions in the three younger classes (kits, yearlings [1.0-1.5-year olds], and subadults [2.0—3.5-year olds]) and grouped all animals > 13.6 kg (4-year olds) as adults. Minitab (Minitab 2000), SigmaStat (Jandel Scientific 1994), and SAS (SAS Institute 1991) were used for all analyses. 7:8:0 8:11:0 5:5:0 10:15:3 1:2:0 2:5:0 12:10:1 14:11:1 NiO) 8:9:2 10:18:4 2:10:0 4:11:0 6:15:0 90:133:11 Total ?: 4d :unknowne 2:3 :unknowne 1:2:0 pheVe) 5:5:0 3:4:0 4:3:0 wine 9:4:1 4:7:4 0:1:0 0:4:0 4:4:1] 0:8:0 2:9:0 3:9:0 39:70:11 # Released Without Transmitters 1:1:0 With Transmitters 2:¢:unknowne 0:1:0 6:6:0 2:3:0 5:7:0 1:2:0 8:8:0 5:7:0 6:11:0 2:1:0 8:6:0 2:2:0 2:2:0 3:6:0 51:63:0 SOONOAAHOOCAOOOCS Unknown Mortality? Results Snares and Hancock traps were used to capture 277 Beaver at 33 locations throughout Wyoming and we eventually transplanted 234 to the 14 release sites. The 43 remaining Beaver either died during trapping (n = 15), transporting (n = 13), or were lac- tating females (n = 15), which we released after cap- ture. Trapping mortality was 10% for Hancock traps and 5.3% for snares, and was not significantly differ- ent (z = -0.07, P = 0.94, df =11). Mortality from traps was due to becoming entangled in snares (n= 11) and being killed by predators while restrained in snares and Hancock traps (n= 4). Trapping success during our five years of trapping was 11.1 trap nights/Beaver or 9.0% (the probability of an individual trap’s capture). Average weight of animals captured was 16.2 kg (range 2.4—31 kg, SD = 6.11) and there was no difference in proportion of males (0.42) or females (0.58) captured (z = 1.23, P = 0.22, df = 122). We transmittered 63 (55.3%) females and 51 (44.7%) males. Of 114 Beaver trapped, equipped with a transmit- ter, and relocated, 34 (30%) died within 180 days (x= 43 days, SD = 37.4) of release (Table 2). Another 7 Beavers (36% total mortality) died prior to the failure of their transmitters (181-503 days). Coyotes (Canis latrans) were responsible for at least 27% of all mortalities, followed by Black Bears (Ursus americanus) (10%), Grizzly Bears (Ursus horribilis) (10%), Mountain Lions (Felis concolor) (2%), and humans (5%). Another 22% (of all mortal- ities) died of unidentified predators and the remain- ing 24% died from undetermined causes. Within 180 days of release, 58 (51%) Beavers either emigrated >10km from the release sites and were not found os Oooo son Oot sOoON OG! Unknown =soocoo7cocoeooooe°ococa Human Predation Mountain Lion otrooococococococjocrtTt Grizzly Bear ooooc*oco=7coomococert Black Bear OnaMeonBOCCOCOCOH= TABLE 2. Fates of introduced Beaver by release location. Letters in () correspond to release sites in Figure 1 and descriptions in Table 1. Coyote ‘Releases at this site failed due to conflicts with irrigation structures and some Beavers were retrapped “Cause of predation was undetermined >Beaver died from undetermined causes ‘Usually kits that could not be sexed (A) Bear Gulch¢ (B) Breteche Creek (C) Bush Creek (D) Currant Creek (E) Deep Creek (F) Ennos Creek (G) Lake Creek (H) Little Red Creek (1) Prairie Dog Creek (J) S. Pine Creek (K) Red Creek (L) Spring Creek (M) Trabing Creek (N) Trout Creek Total eu== ene a 64 THE CANADIAN FIELD-NATURALIST again, or developed faulty transmitters, which made them impossible to relocate. The Kaplan-Meier survival estimates for all Beavers were 0.49 for 180 days and 0.433 for 360 days and did not differ between age classes (Table 3). Beavers that died lived for an average of 86 days (range 1-503 days, SD = 114.8) until death, however eight died within seven days of release. All Beavers (that we found), except one, died within 0.5 km of the release site (the exception was found 0.75 km upstream of the release site). Twenty-three (19%) Beavers lived > 180 days and eventually built dams and lodges in the drainages where they were released. Additionally, a minimum of 10 (actual number could not be determined since we did not retrap Beavers after release) of the other 120 Beavers released without transmitters were also found with dams and lodges within 3 km of the release sites (some of these may have been Beaver with faulty transmitters, although we feel this was unlikely). We released an average of 17 Beaver at each of our sites in an attempt to get animals to establish. Beaver successfully established at 13 of the 14 release sites and, as of September 2001, the 13 sites were still occupied. At the unsuccessful site Beavers had conflicts with irrigation structures (e.g., damming irrigation ditches) and they were removed. We were unable to identify any variables in our analyses that significantly influenced the probability of success for beaver relocations. P values were > 0.2 and R-squared values were <0.10 for all vari- ables and models tested. The 2—3.5 year old age class had greater average occupancy at a site (Figure 2), although this relationship was not significant (P= 0.225, df 3). All kits and yealings (n = 12) either died (n=5) or emigrated (n= 7) from the release site prior to 108 days and none were observed constructing dams and lodges. Discussion Limited range of transmitters and transmitter fail- ure may have influenced the number of animals we Vol. 116 - found after release (Rothmeyer et al. 2002), subse- quently increasing the number of Beavers assumed to have emigrated. Advertised range of the internal transmitters was 0.5 km but we found that the range was usually limited to 200 m, and animals within dens were not located until we were within 50 m of the transmitter. Range of the tail-mount transmitters was better but never approached the 1-km advertised range. Walking up and downstream of the release site within 20 m of the creek bed was necessary to determine movements and mortality. Animals emigrating or not found after release may have had higher survival rates but we believe this is improbable. More likely, these Beaver were killed and cached in holes and dens, or dragged out of range of receivers. Animals moving out of the vicini- ty of the release area may also have experienced higher predation rates due to increased exposure time and less time spent hiding in dense vegetation or constructing dens. Beavers, sympatric with Black Bears on islands in Lake Superior traveled shorter distances from water than those found on islands where bears were not present, posssibly a direct attempt to avoid predation (Smith et al. 1994). Survival estimates for unexploited (i.e. untrapped) adult beaver are generally 0.80 (Boyce 1974; Bergerud and Miller 1977; Bishir et al. 1983) but these animals have ponds and lodges for escape. High mortality (5/10) and loss (4/10) (never relocat- ed) rates were reported for beaver translocated in the James Bay area of Quebec, Canada (Courcelles and Nault 1983). Translocated animals undoubtedly have a higher susceptibility to predation for many reasons including unfamiliarity with the surrounding habitat, reduced fitness due to trapping stress, and possible exposure to more numerous and greater varieties of predators (Griffith et al. 1989; Stanley-Price 1989). Beaver have many natural predators and do not avoid predation through fighting, preferring instead to use water as an escape medium (as reviewed in Smith et al. 1994). Without ponds and dens to use for escape, Beaver are vulnerable to predation. TABLE 3. Kaplan-Meier 180 and 360 day survival estimates, sample sizes, 95% CIs, and SEs for Beavers translocated in Wyoming. Survival estimates were not computed for Beaver < 2 years old since 100% either died or emigrated prior to 180 days. Age N n n n Survival Class Total Surviving Dying Unknown Estimate 95% CI SE 180 days All ages 114 23 34 58 0.493 0.358-0.628 0.068 2.5-3.5 years old 52 13 20 19 0.436 0.271-0.602 0.085 4+ years old 51 10 8 33 0.658 0.449-0.868 0.107 360 days : All ages 114 13 36 66 0.433 0.268-0.598 0.084 2.5-3.5 years old 52 fi 22 23 0.353 0.167-0.538 0.095 4+ years old 51 6 8 S. 0.658 0.449-0.868 0.107 4animals either emigrated > 5 km or transmitters failed 2002 MCKINSTRY AND ANDERSON: TRANSPLANTED BEAVERS IN WYOMING 65 600 500 » io) (jo) 300 200 Success (days) = [=) oO Kits Yearlings Subadults in Adults (1.5 yrs old) 3.5 yrs old) (> 3.5 yrs old) Age Class FIGURE 2. Average survival, in days (n), for four age classes of relocated beavers. Error bars depict 95% CI’s. Naturally dispersing Beaver have been seen in upland areas > 1 km from water sources and have been observed crossing mountain passes (Smith 1980) far from water where they would be extremely vulnerable to predation. Movements across upland areas are undoubtedly successful but we question the frequency of this success in areas like Wyoming where multiple predators occur sympatrically. Translocated Beaver in North Dakota, Wisconsin, and Maine moved an average of 14.6, 7.4 and 11.2 stream km, respectively, although the longest move- ment was 238 km (Hibbard 1958; Knudsen and Hale 1965; Hill 1982) and naturally dispersing Beaver in Idaho and Quebec moved an average 8.5 and 18 km (Leege 1968; Courcelles and Nault 1983). While movements greater than 3 km are common for Beaver, exposure rates certainly increase during long movements through non-ponded habitat. Our ani- mals may have been attempting to return to trapping locations, which were greater than 160 km away. Since our goal was to establish Beavers in uncolo- nized areas, predation risk was unavoidable and high predation and emigration losses should be consid- ered when planning releases. Wildlife managers in Europe have made several attempts to transplant Beaver (C. fiber) with varied success (Zurowski and Kasperczyk 1988; Hartman 1994; Nolet and Baveco 1996). Initially, mortality and emigration rates were high (14-36% and 23%, respectively) and were greatest in juvenile and year- ling animals. Mortality rates declined with the estab- lishment of dens and dams and populations became well established over time. However, transplanting in Europe usually occurs in major river systems where flooding of the dens is the primary disruptive event and predator populations are not as large (Nolet and Baveco 1996). In Wyoming, predator communities are well developed and are thought to be increasing (McKinstry and Anderson in review). Additionally, Beaver introduced in large rivers (> 2.83 m/sec) have greater aquatic escape cover than Beaver released in the small (< 0.283 m/sec) streams where we were working. The value of riparian areas for wildlife has been emphasized by many authors (as reviewed by Naiman et al. 1988), but predator-prey relationships in riparian areas and how they relate to habitat quality and availability are not well understood (Smith et al. 1994). During drought periods animals may concen- trate in riparian areas along with their respective predators. These predators, while normally dependent upon another prey species, may find Beaver easy prey. Many wildlife species use riparian areas more frequently during fall due to increased forage and water in the riparian areas. Predators may also con- centrate in these areas to take advantage of higher prey densities, greater water availability, and lowered temperatures. Our releases were primarily (71%) done during the fall (15 August—10 October) to take advantage of the Beaver’s natural tendency to begin construction of dams and lodges in the fall (Vore 1993). We did spring releases to establish Beaver in 66 THE CANADIAN FIELD-NATURALIST ephemeral streams that were normally dry by mid August and to supplement one or two Beaver that had become established the year before. Relocations in the spring may have lower predation risks, although we did not see this relationship in our analyses. Several biologists suggested that we create small ponds at release sites to provide temporary refuge for released Beaver. In the two instances (Bear Gulch and Spring Creek; see Table 2 for specific numbers released) where we released Beaver into remnant ponds they emigrated from those areas within 10 days and constructed their own dams and lodges elsewhere in the drainage, although the remnant ponds may have provided them with initial protec- tion from predators and helped to acclimate them to the area. Furthermore,. creating small ponds is cost prohibitive and unlikely to be used in future trans- plants, therefore we did not consider this as a legiti- mate tool in our transplants. Our goal was to establish Beaver at release sites, therefore we continued to release Beaver until dams and lodges were constructed. On average we released 17 Beaver/site before they constructed a dam and lodge and successfully reproduced. In 11 instances a single Beaver created a pond with a lodge and we transplanted additional animals in the hope that they would pair-up. We expected that many Beaver would emigrate from our release sites in efforts to return home, search for mates, or look for more suitable habitat. High predation and emi- gration rates for introduced Beaver should be expect- ed (Griffith et al. 1989) and planned for in any Beaver translocation project. We found no significant predictors of success or survival in our analyses. The 2—2.5 year-old Beavers had greater average success (Figure 2) and other researchers (Vore 1993; P. Jensen, Pennsylvania State University, personal communication) have sug- gested that this age class may be more suitable for recolonizing new areas since they are predisposed to emigrating, establishing new territories, constructing dams and lodges, and attracting mates. Our results are inconclusive but we recommend translocating Beavers > 2 years old since mortality and emigration of our younger animals totaled 100% within six months. Our results also call attention to programs where only one or two Beaver are relocated to either unoc- cupied habitat, usually under the pretext of improv- ing habitat, or to areas with existing Beaver popula- tions. These activities are common in the western US and are usually carried out by district biologists or conservation officers to remove nuisance Beaver without using lethal techniques. We suggest that’ unless managers are committed to successfully intro- ducing Beaver through planned introductions, moni- toring, and follow-up releases their time and money may be better spent on other duties. Vol. 116 - In summary, we found that Beaver could be used to create natural wetlands and improve riparian habi- tat in 13/14 streams where we relocated them. Mortality (30%) and emigration (51%) totaled 81% within the first 6 months of release and we needed to relocate an average of 17 Beaver/site to get a pair to establish and reproduce. Additional releases were also needed to augment single animals that had become established. Wildlife response to our created habitats was immediate (McKinstry et al. 2001) and landmanagers found that the habitats were valuable to both wildlife and livestock. We caution that releases should only be used in drainages where con- flicts with irrigation or road crossing structures are minimal and preferably where the drainage is con- trolled by a few landowners to simplify evaluation and management. Acknowledgments This research was supported by grants from the Wyoming Game and Fish Department, U.S. Fish and Wildlife Service, Rocky Mountain Elk Foundation, Ducks Unlimited, National Rifle Association, National Fish and Wildlife Foundation, North American Wetlands Conservation Council, Jack’s Plastic Welding, George E. Menkins Memorial Fellowship, and the University of Wyoming. Field assistance, provided by Rory Karhu, E. Hardgrave, S. Rothmeyer, S. Mohren, and K. Rompola was much appreciated. Our thanks are also extended to the Breteche Creek Foundation, T. Malmberg, M. Miller, U.S. Forest Service, US Bureau of Land Management, the State of Wyoming, and numerous other landowners for allowing us access to release sites. L. Apple, J. Vore, K. Gordon, G. Butler, and two anonymous reviewers suggested revisions that improved the manuscript. Literature Cited Apple, L. L., B. H. Smith, J. D. Dunder, and B. W. Baker. 1985. The use of Beavers for riparian/aquatic habitat restoration of cold desert, gully cut stream sys- tems in southwestern Wyoming. Pages 123-130 in Investigations on Beavers. Edited by G. Pilleri. Institute of Brain Anatomy, Berne, Switzerland. Barnes, W. J., and E. Dibble. 1986. The effects of Beaver in riverbank forest succession. Canadian Journal of Botany 66: 40-44. Bergerud, A. T., and D. Miller. 1977. Population dynam- ics of Newfoundland Beaver. Canadian Journal Zoology 55: 1480-1492. Bishir, J.,. R. A. Lancia, and H. E. 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Acta Theriologica 33: 325-338. Received 1 November 2000 Accepted 18 March 2002 Songbird Community Composition Versus Forest Rotation Age in Saskatchewan Boreal Mixedwood Forest END E. Cumminc!, and ANTONY W. DIAMOND? '1542 Empress Avenue, Saskatoon, Saskatchewan S7K 3G3 Canada Atlantic Co-operative Wildlife Ecology Research Network, University of New Brunswick, P.O. Box 45111, Fredericton, New Brunswick E3B 6E1 Canada Cumming, Enid E., and Antony W. Diamond. 2002. Songbird community composition verses forest rotation age in Saskatchewan boreal mixedwood forest..Canadian Field-Naturalist 116(1): 69-75. Songbird communities were characterised using unlimited-distance point counts in four age-classes of boreal mixedwood forest in central Saskatchewan in 1991. Forest age-classes surveyed were: mature stands (50-60 years old), rotation age stands (80-90 years), old stands (100-110 years), and very old stands (>140 years). Ten species (Winter Wren, Troglodytes troglodytes; Golden-crowned Kinglet, Regulus satrapa; Ruby-crowned Kinglet, R. calendula; Swainson’s Thrush, Catharus ustulatus; Tennessee Warbler, Vermivora peregrina; Blackburnian Warbler, Dendroica fusca; Magnolia Warbler, D. magnolia; Bay-breasted Warbler, D. castanea; Rose-breasted Grosbeak, Pheucticus ludovicianus and Evening Grosbeak, Coccothraustes vespertinus, were significantly more abundant in forest that was older than rotation age. Four of these species (Winter Wren, Tennessee, Magnolia and Bay-breasted warblers), reached their highest densities only in the oldest (>140) age class. Two species (Red-eyed Vireo, Vireo olivaceus, and Ovenbird, Seiurus aurocapillus) were signifi- cantly more abundant in forest that was rotation age or younger. In boreal mixedwood forest, habitat-selection patterns of songbirds suggest that many species, not also occurring in rotation-aged or younger stands, may require substantially older forest in order to persist. Key Words: songbirds, boreal, mixedwood forest, rotation age. Boreal forest is one of the largest terrestrial ecosys- | comes under increasing pressure from harvesting, tems on the planet (Shugart et al. 1992; Apps et al. foresters will be expected to maintain biodiversity 1995). It comprises 20% of the world’s forest and is (CCFM 1992*; Freedman et al. 1994; Middleton the largest terrestrial ecozone in Canada, covering 1994*) but currently lack the information required to more than one-third of the country (Rowe 1972; doso. When a previously little-exploited segment of Shugart et al. 1992). The southern part of this forest, the forest resource, such as old-growth mixedwood the boreal mixedwood, supports the best tree growth _ boreal stands, becomes the focus of intensified har- (Rowe 1972) and is under the most pressure from __ vesting, the assessment of impacts on biodiversity human activities. In the three Prairie Provinces, more becomes an urgent priority. than 90% of the boreal mixedwood forest has been Our study was designed to address the question: leased for harvesting under Forest Management “Are there elements of the biological community of Agreements (Cummings et al. 1994; Pratt and Urqhart old-growth boreal mixedwood forest whose survival 1994; Alberta Environmental Protection 1998*). In may be threatened by harvesting?”. In his 1977 study Saskatchewan, over 80% of trees harvested come on boreal songbirds, Erskine suggested that the from mixedwood forest, predominantly White Spruce “...mature stage of the most fertile needle-leafed (Picea glauca) and Trembling Aspen (Populus tremu- community” is biologically the most diverse commu- loides), especially from the older age classes (Spencer __ nity in the boreal forest and “...deserves more atten- 1993). This type and age of forest accounts for, at tion from environmental impact studies than they most, 10% of the commercial forest (Kabzems et al. have received’. Specifically, we used songbirds as 1986), yet it is also the most biologically diverse habi- focal organisms, expecting that they might be useful tat in the region (Welsh 1981; Harris et al. 1984; indicators of impacts on other organisms, and per- Acton et al. 1998). Thus, the biologically richest haps of biodiversity as a whole (Freedman et al. stands are being harvested in a proportion far exceed- 1994). Songbirds are primarily insectivorous, feed- ing their abundance (Erskine 1977). ing sufficiently close to the top of their food-chains Canada is required under the Biodiversity Con- to be potentially useful indicators of impacts on a vention to ensure the ecological sustainability of all wide range of organisms (Diamond and Filion 1987; its ecosystems (Biodiversity Science Assessment Furness and Greenwood 1993). Team 1994*), and the global market is demanding We chose a range of stand ages around the pro- that suppliers of forest products demonstrate the posed rotation age (80—90 years), to assess whether, environmental sustainability of their harvesting prac- _in the extreme case of all stands older than rotation tices (Canada 1995*). As boreal mixedwood forest age being eliminated from the landscape, an impact 69 oe. ee ee eee ——— eee ee ee 70 THE CANADIAN FIELD-NATURALIST Vol. 116 - TABLE 1. Species which occurred significantly more or less frequently in a given forest age (Anova with Tukey’s test). Species E Winter Wren 87.1 Tennessee Warbler 3.0 Magnolia Warbler 4.8 Bay-breasted Warbler 6.9 Ruby-crowned Kinglet 6.1 Blackburnian Warbler El Black-throated Green Warbler 4.3 Rose-breasted Grosbeak 4.5 Swainson’s Thrush he Red-eyed Vireo 4.3 Ovenbird 12.6 7<0.05, 770.01, **t<0.001 on the bird community could be expected. Our youngest age-class was 30 years younger than rota- tion age. We did not study stands younger than this, because other studies have shown the songbird com- munity to be very different between young (<30) and old (>80) forest (Titterington et al. 1979; Schiek et al. 1995; Hobson and Bayne 1999). In addition, intensified management will lead to an increased abundance of younger age-classes at the expense of those older than rotation age (Spencer 1993; Weyerhauser Canada 1998*). It is already known that a reduction in stand age may have negative effects on some bird species (Titterington et al. 1979; Welsh 1981; Farr 1993; Schiek et al. 1995), but effects on particular species in spruce-dominated mixedwood forest have not pre- viously been addressed. Previous studies have shown that old mixedwood forests have a rich assemblage of bird species (Erskine 1977; Kirk et al. 1996). Many of these species do not occur in younger forest, or in hardwood dominated forest, or do so only at low den- sities (Stelfox 1995; Hobson and Bayne 1999). Methods This study was carried out in summer 1991 in and around Prince Albert National Park, Saskatchewan (53° 35'N 106° 00'W) (see Bayne and Hobson 1997), in boreal mixedwood forest (Kabzems et al. 1986; Acton et al. 1998). Ten study sites were established in forest of four different ages: three stands >140 years old, three 100-110 years old, two 80-90 years old and two 50-60 years old, and were aged by coring six of the largest trees on each site. All sites were mixed- wood forest and were as similar as field conditions allowed. Sites were pre-selected for similarity using Saskatchewan forest inventory maps, which are based on similarity of tree density and species composition (Kabzems et al 1986). Ground truthing showed stands _ were similar in species composition and tree density, however, older sites did tend toward greater abun- dance of conifers (Cumming 1995). The largest and most abundant trees in all stands, were White Spruce DF P Significant Age 3 ae 140 > 3 i 140 > 3 id 140> 3 aR 140> 5 -” 100 > S: or 100 > 3 a 100 > S er 100 > 3 ai 50 < 3 a 80> 5 ee 140< and Trembling Aspen. Two other species, White Birch (Betula papyrifera) and Balsam Fir (Abies bal- samea) also occurred (Cumming 1995). The main shrubby species were young Balsam Fir, White Birch, Alder (Alnus sp.) and Beaked Hazel (Corylus cornu- ta). Ground cover was mainly composed of several species of moss; Bunchberry (Cornus canadensis); Twinflower (Linnaea borealis) and Wild Sarsaparilla (Aralia nudicaulis). Birds were counted using the unlimited-distance point count method (Bibby et al. 1992), which is widely used as the method of choice when large areas need to be surveyed rapidly and efficiently (Verner 1985; Farr 1993; Bancroft et al. 1995). The method is a good indicator of relative abundance (Verner and Ritter 1985; Bancroft et al. 1995) and an efficient method for comparing species composition and richness between habitats (Blondel et al. 1981). Point counts were at least 200 m apart and 100 m from a stand edge; these distances minimise the chances of double-counting, and including species from other habitats, respectively. Surveys were con- ducted between 04:00 and 09:00, by the same observer (EEC), and included all birds seen or heard in 10 minutes. Each site was surveyed twice during the breeding season; early (1-14 June) and late (20 June—4 July). Such spacing of surveys gives a more representative picture of a bird community than either one survey, or two carried out on consec- utive days (Hilden 1981; Skirven 1981). Because sampling effort differed between stands, (five point count stations/site in the oldest category, and three/site in the other three categories), the data were standardised by dividing the maximum number of individuals of each species recorded in a particu- lar age class by the number of point counts. This gives the probability of detection for a particular species in that age of forest. Birds that were flying overhead, or had very large territories, such as rap- tors, ravens, and woodpeckers were not included in the analysis. 2002 CUMMING AND DIAMOND: SONGBIRD COMMUNITY COMPOSITION 71 Forest Age (years) cee me <().20) —— 0.2 to O49 0.50 to 0.79 mmm () §() to 1.00 a > 1.00 Bird Species 50-60 80-90 100-110 140+ Winter Wren es - Cape May Warbler Western Tanager White-winged Crossbill Gray Jay Golden-crowned Kinglet Rose-breasted Grosbeak Evening Grosbeak Brown Creeper Boreal Chickadee Ruby-crowned Kinglet Red-breasted Nuthatch Solitary Vireo Yellow-rumped Warbler Mourning Warbler Chipping Sparrow White-throated Sparrow Pine Siskin Swainson's Thrush Tennessee Warbler Blackburnian Warbler Black-throated Green Warbler Magnolia Warbler Bay-breasted Warbler Red-eyed Vireo Ovenbird —_ oo Number of Species 15 4 more abundant in post rotation age sites (p<0.05) ok *k *K *K i) i) i) ON more abundant in sites at or below rotation age (p<0.05) FiGurE 1. Probability of detecting a given species of bird at a point count in the different forest age classes. Vegetation was measured in 0.04 ha circles (22 metres in diameter), using a modified James and Shugart (1970) method. Two such circles were locat- ed near each point count station, one centred on the point count location, with the other located in a ran- dom direction 50 metres away. In each plot, the species, diameter (dbh), and height of each tree was recorded. Plants with a dbh greater than 7 cm were considered trees, while those less than 7 cm dbh were Classified as shrubs (James and Shugart 1970). Percent shrub cover was estimated for the entire plot, and ground cover was estimated in four randomly located 1m? quadrats. Data were analysed using l-way Anova (Zar 1996) using the statistical package SPSS. In the case where more than two categories were used, a Tukey’s HSD test was applied to find where differ- ences had occurred (Zar 1996). Both bird and vege- tation data were analysed using four age classes and by pooling data for two age classes, either post-rota- tion age or rotation age and younger. Results The number of songbird species recorded in each habitat progressively increased with age, from youngest to oldest the numbers of species were; 15, aula ee eee ae —— wee ewe - ee ee eee v2 TABLE 2. Species which are found significantly more often in post-rotation age forest. Species F DF P Winter Wren 15.8 1 ae Golden-crowned Kinglet Se) * Ruby-crowned Kinglet 5.0 1 a Swainson’s Thrush 533 ] % Tennessee Warbler 4.1 l ¢ Magnolia Warbler 6.1 1 5 Blackburnian Warbler 8.7 1 ee Bay-breasted Warbler 39 1 v5 Rose-breasted Grosbeak 4.7 1 x Evening Grosbeak iG 1 ae Red-eyed Vireo’ 6.8 1 at 1 Ovenbird" 7a F<),05, **#140 year old forest (Table 1). Ruby-crowned Kinglet, Rose-breasted Grosbeak, Blackburnian and Black-throated Green warblers (Dendroica virens) were most abundant in the 100-110 year old forest while Red-eyed Vireos were most abundant in the 80-90 year old forest. Swainson’s Thrushes were least abundant in the 50-60 year old forest, and Ovenbirds were least abundant in the >140 year old forest (Table 1). Pooling the data into pre and post- rotation age showed ten species to be significantly more abundant in post-rotation aged forest (Table 2). Only Ovenbird and Red-eyed Vireo were more abundant in forest of rotation age and youncer (Table 2). The percent of coniferous trees relative to decidu- ous trees increased with increasing forest age (Table 3). The average height of the forest was not signifi- cantly different between the three oldest ages, but THE CANADIAN FIELD-NATURALIST Vol. 116 - the average tree height of the 50-60 year old forest was significantly shorter than the other three (Table 3). The amount of shrub cover and moss cover was significantly higher in the oldest forest, while the amount of litter was significantly higher in the youngest forest (Table 3). The average height of the combined post-rotation age forest (20.4 + 7.0 m) was significantly taller than the rotation age and younger forest (15:8 + 3.9 m) (Fo = 300 P< 0.001). | Discussion All bird species found in the youngest stands also occurred in older forest; however, the reverse was not true. Species were added to the forest with increasing age, and species richness increased with stand age. We sampled more old than young stands, therefore, with increased sample effort some of the missing species may indeed be detected in younger forest. However, our data indicate a definite trend for higher species richness in forest that is older than rotation age and for many species there is also a trend for greater abundance in forest older than rotation age. This may be caused by old forest hav- ing higher habitat heterogeneity than young forest, which is thought to contribute to its increased species richness ( Meslow et al. 1981; Hunter 1990). This increased heterogeneity has a variety of causes, including windthrow gaps, and tree disease and death, which allows for infilling with various ages and species of trees (Erskine 1977; Hunter 1990). Studies in Maine, Ontario, and Alberta found Ruby and Golden-crowned kinglets, Winter Wrens, crossbills (Loxia spp.), Cape May, Blackburnian, and Bay-breasted warblers were more abundant in mature to old forest and present at either much lower densities, or absent, in younger stands (Titterington et al. 1979; Welsh 1981; Schieck et al. 1995). A small sample size was the most likely reason we did not find Cape May Warbler, Western Tanager (Piranga ludoviciana) and White-winged Crossbill (Loxia leucoptera) significantly correlated with post- rotation aged forest. However, as they were observed only in the oldest forest sampled, this suggests a TABLE 3. Percent of coniferous and deciduous tree cover and average height at different ages, and percent of shrub and ground cover at different ages. Percentages are mean + | standard deviation. Tall Shrub % % Age Conifer Decid. Height (m) % Cover 140+ 80 20 20:1 47.5 41.7 + 24.6* 100-110 65 35 21.1 26.1 28.8 + 22.3 80-90 48 op) 18.3+4.3 22.9 2:38.60 50-60 49 51 13.9 + 2.0* phe ee Ground Cover % Moss % Herb % Litter % Grass 53.0 + 28.3* 4.3+3.5 42.8 + 27.2 0 9.4+ 17.4 22.5+16.5 65.6+ 20.9 2,28 Tal 2.5 + 5.0 25.0+28.9 70.0+29.4 2.5 +£5.0 3.1+2.4 44+2.3 92.5 + 12.9* 0 *Indicates means that are significantly different at P<0.05;1-way Anova;Tukey’s HSD. 2002 trend for these species to use mature to old forest, especially as they have been found to be significant- ly associated with older forest in other studies (Farr 1993: Benkman, 1993). Some species did not show a significant difference in their habitat preference and occurred at similar densities in all ages tested. Species such as, Solitary Vireo (Vireo solitarius); Yellow-rumped Warbler (Dendroica coronata); Chipping Sparrow (Spizella passerina) and Pine Siskin (Carduelis pinus), have been found to be associated with coniferous forest, including Jack Pine (Pinus banksiana), rather than forest age per se. (Kirk et al. 1996); and species such as White-throated Sparrow (Zonotrichia albicolliss), are widely distributed across a great many forest types and ages (Kirk et al. 1996; Hobson and Bayne 1999). Many of the 26 species sampled showed some degree of dependence on forest as old as, or older than, the rotation age of 80-90 years. Eight species (30% of the total community) occurred exclusively in stands substantially older than rotation age, and three more occurred only in these stands or those of rotation age; thus 42% of the species in this commu- nity showed a marked degree of dependence on mature or old stands. In addition, five other species (Swainson’s Thrush, Tennessee, Blackburnian, Magnolia, and Bay-breasted warblers), that occurred in all age classes, were significantly more abundant in the post-rotation age forest. We conclude that 62% of these species (16/26) are likely to suffer some reduction in population density if stands older than rotation age are altogether lost from the land- scape. Several of the species which occurred in younger stands did so at low abundances; for these species, young stands could potentially represent population sinks (van Horne 1983; Pulliam 1988), in which populations can only be maintained by immigration. Many of the species that were age-sensitive also appear to be area-sensitive (Hobson and Bayne 2000). As these species appear to show sensitivity to age and fragmentation, we suggest that there is a need for further research on the habitat requirements and reproductive biology of these species. This is especially true as many of these species may also be facing habitat loss in the southern fringe of the bore- al mixedwood forest (Cumming et al. 2001; Hobson et al. in press). We conclude that managed boreal mixedwood forest should maintain some stands of at least 120 years or older if the communities characteristic of the whole forest ecosystem are to be sustained. Natural disturbance patterns could be used to guide the proportion of the landscape to be retained in this age-class. The question of the most appropriate size and configuration of such old-growth stands, in relation to conservation of songbirds, was not CUMMING AND DIAMOND: SONGBIRD COMMUNITY COMPOSITION qs addressed by our study, but we draw attention to abundant evidence that reduction in forest age, and fragmentation of forest patches (a process that has accelerated in central Saskatchewan in recent years, Fitzsimmons et al. 1997*) can have serious impli- cations for the sustainability of songbird popula- tions (Schieck et al. 1995; Villard et al 1995; Cumming et al. 2001) Acknowledgments This study was carried out in partial fulfilment of the requirements for the M.Sc degree at the University of Saskatchewan by E. E. Cumming. Parks Canada and Weyerhauser Canada Ltd. allowed field work to be carried out in Prince National Park and on Weyerhauser lease, respectively. Funding for this project was provided by the Canadian Wildlife Service (which also provided equipment and office facilities at the Prairie and Northern Wildlife Research Centre), Prince Albert National Park, Nature Saskatchewan and the Saskatchewan Wildlife Habitat Development Fund. This manuscript was improved by helpful comments by A. J. Erskine, K. A. Hobson, S. L. Van Wilgenburg and an anony- mous reviewer. Documents Cited (marked * in text) Alberta Environmental Protection. 1998. The Final Frontier: Protecting landscape and biological diversity within Alberta’s Boreal Forest Natural Region. Report for the Special Places 2000 Provincial Co-ordinating Committee. Natural Heritage and Evaluation Branch. March 1998. Protected areas report number 13. Biodiversity Science Assessment Team. 1994. Biodi- versity in Canada: a Science Assessment. Minister of Supply and Services. Ottawa. Canada. 1995. The state of Canada’s forests 1994. Minister of Supply and Services. Ottawa. Canadian Council of Forest Ministers (CCFM). 1992. Sustainable forests — a Canadian commitment. Minister of Supply and Services Canada, Ottawa. Fitzsimmons, M, L. Patino, P. MacTavish and P. Farrington. 1997. Estimating changes in forest area in central Saskatchewan, Canada. Poster, presented at the 3rd National Ecological Monitoring and Assessment Network Conference, Saskatoon, Saskatchewan, Feb- ruary, 1997. Middleton, J. 1994. Effects of forestry on biodiversity in Canada. Pages 51-58 in Biodiversity Science Assess- ment Team. Biodiversity in Canada: a Science Assess- ment. Minister of Supply and Services. Ottawa. Weyerhauser Canada. 1998. Twenty-year forest manage- ment plan and environmental impact statement. Weyerhauser Canada and Saskatchewan Environment and Resource Management. Prince Albert, Saskatchewan. Literature Cited Acton, D. F., G. A. Padbury, and C. T. Stushnoff. 1998. The ecoregions of Saskatchewan. 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Impact on bird populations of harvest- ing in the boreal mixedwood forest. Pages 155-167 in Boreal Mixedwood Symposium. Edited by R.D. Whitney and K. M. McClain. Ontario Ministry of Natural Resources. Zar, J. H. 1996. Biostatistical Analysis. Third edition. Prentice-Hall Inc., Upper Saddle River, New Jersey. Received 6 November 2000 Accepted 19 March 2002 Status of Redside Dace, Clinostomus elongatus, in the Lynde and Pringle Creek Watersheds of Lake Ontario JEFF J. ANDERSEN Central Lake Ontario Conservation Authority, 100 Whiting Avenue, Oshawa, Ontario L1H 3T3 Canada Andersen, Jeff J. 2002. Status of Redside Dace, Clinostomus elongatus, in the Lynde and Pringle creek watersheds of Lake Ontario. Canadian Field-Naturalist 116(1): 76-80. Populations of the Redside Dace, Clinostomus elongatus, are declining throughout most of their Ontario range. Since 1959, Redside Dace have been found at 20 sampling sites in Lynde and Pringle creeks within the Town of Whitby, Ontario. These collection sites were reassessed in 1999 and 2000 to investigate the status of the Redside Dace within these systems. A total of 20 Redside Dace were captured at four of the 20 sites. Populations are now either severely limit- ed or absent from Pringle Creek and are now confined to upstream areas in Lynde Creek. Redside Dace were confined to 24 and 34 order coolwater streams with variable substrata, and these habitats displayed a 100% canopy closure by grass species. Potential Redside Dace habitats are identified within the watershed to provide general guidelines for future land development. Key Words: Redside Dace, Clinostomus elongatus, fish, minnows, threatened species, special concern, vulnerable species. The Redside Dace has a discontinuous global dis- reported to be quite sensitive to turbidity (Scott and tribution and is found only in North America. In the Crossman 1973; Trautman 1981). Redside Dace dif- western portion of its range, it occurs in the upper fer from other members of the family Cyprinidae in Mississippi and Lake Michigan drainages, and, inthe that approximately 77% of their diet consists of eastern portion, in the Lake Erie, Lake Huron, Lake insects. Daniels and Wisniewski (1994) suggest a Ontario and upper Susquehanna River drainages. dominance for flying insects made up mainly of Page and Burr (1991) describe the Redside Dace as__danceflies, caddisflies and midges. The Redside locally common in the eastern part of its range Dace possesses large eyes and has a very large although it is declining in many areas. The Redside mouth which allows it to catch these in flight by Dace is localized and rare in the western part of its leaping out of the water. range. Collection records for Redside Dace in Lynde and Pringle creeks, located in the Township Ontario indicate “the general distribution included of Whitby, Region of Durham, Ontario, centered at tributaries of Lake Ontario from Lynde Creek in the 43° 56’ 30” W, 78° 59’ 10” N (Figure 1), are the only east to Spencer Creek on the west, and north to the creeks reported to support Redside Dace within the headwaters of these systems” and “in the Lake Central Lake Ontario Conservation Authority Simcoe drainage limited to Kettleby Creek” (Parker (CLOCA) watershed (CLOCA 1996—2000*). et al. 1988). They are also known to occur in the Redside Dace habitat was described by Parker et Saugeen River watershed, Gully Creek and Two Tree al. (1988) as pools and slow moving sections of rela- River in the Lake Huron Drainage, and from Irvine _ tively small headwater streams which have a pool rif- Creek in the Lake Erie Drainage. fle habitat. Stream sections with overhanging bushes The Redside Dace, Clinostomus elongatus, was and herbaceous plants to provide some cover were classified as Vulnerable in Canada by the Committee found to be particularly suitable. Bottom substrates on the Status of Endangered Wildlife in Canada ___ were usually composed of boulders, rocks gravel or (COSEWIC) in 1987. In 2000, COSEWIC changed _ sand, often with a shallow surface covering of detri- the “Vulnerable” classification to Special Concern _ tus and silt. Streams were clear and colourless in con- (COSEWIC 2000). Studies are currently underway junction with hard substrates and clear to brown to reassess the status of the Redside Dace on behalf tinged streams with organic substrates. This species of COSEWIC (E. Holm, Royal Ontario Museum, prefers clear water and is quite sensitive to turbidity. personal communications January 1999 to January Novinger and Coon (1998) describe a micro-habitat 2000). The Ontario counterpart of COSEWIC, the preference for mid-water positions in the deepest Committee on the Status of Species at Risk in _ parts of pools under overhanging vegetation. Ontario (COSSARO) designated the Redside Dace The status of the Redside Dace in Lynde and as Threatened in Ontario in May 2000 (COSSARO Pringle creeks had not been assessed since 1985. 2000). * Due to the stresses of increased development within Redside Dace prefer cool, clear, flowing waters the Lynde and Pringle creek watersheds and the sen- with a substrate composed of gravel or stone and are _ sitivity of Redside Dace to increased turbidity, an 76 7. | 2002 ANDERSEN: REDSIDE DACE IN LYNDE AND PRINGLE CREEK WATERSHEDS TF Legend = Potential] Habitat based on infrared Piotegrapty Interpretation * = flistorical and Conicmporary Capture > Historical Capture Only = = Waicrshed Boundaries ——- = Roads { Map Not to Seuic 7 Pringle Creek Watershed FiGureE 1. Historical (@) and contemporary (%*) Redside Dace, Clinostomus elongatus, collection sites and potential habitat on Lynde and Pringle creeks. impact on the Redside Dace population would appear inevitable. The purpose of this study would be an attempt to summarize historical capture data, resurvey all known capture sites, identify the current status and range of known Redside Dace habitat and predict potential Redside Dace habitat within the Lynde and Pringle creek watersheds. Methods Historical Capture Mapping All historic collection sites from 1959 to 1999 were taken from the CLOCA Fisheries Database and mapped on a 1:10000 Ontario Base Map (Figure 1). Exact locations were difficult to assess due to location reporting in previous studies which were given in gen- eral or relative terms. Survey locations were, there- fore, derived from a combination of the above infor- mation, landmarks denoted in field records as well as conversations with individuals who conducted the studies (E. Holm, personal communication 1999). Collection Sample collections for the historical studies on the Lynde and Pringle were conducted with seine nets and dip nets only (Archives of Ontario 1959* and Tumey 1984*) or a combination of seine and elec- trofishing (Holm and Crossman 1986*), and were qualitative in nature. For the current study, qualita- tive fish samples were obtained using the single-pass electrofishing method described in Stanfield et al. (1997). Sampling areas coincided with areas of his- toric Redside Dace occurrence. This method was effective and least harmful to the fish compared to netting methods due to the speed of the sample and the relatively short electrical field exposure time. The length of steam electrofished ranged from 18 to 123 metres and averaged 49 metres. The electrofish- er was set at 45 Hertz at 4 seconds with an output of 300 Volts. Field collections were conducted July and September 1999 and August and October 2000. All species captured were noted and Redside Dace cap- ture locations were determined using UTM coordi- nates. All species were identified in the field and a sample of Redside Dace specimens from each cap- ture site were sent to the Royal Ontario Museum for verification. Habitat Mapping Potential habitat was identified using Ontario Ministry of Natural Resources rectified colour infrared photographs (Ontario Ministry of Natural Resources 1997). Areas were identified that were 2nd, 3rd and 4th order streams on the Lynde and 78 THE CANADIAN FIELD-NATURALIST Pringle creeks. These systems were within agricul- tural areas with riparian vegetation made up of main- ly grass species. Results Historical Data Summary Historically, Redside Dace have been noted in both Lynde and Pringle creeks located within the Town of Whitby (Figure 1). In four surveys on the Lynde and Pringle creeks the capture of the Redside Dace was reported (see site codes in Table | and Figure 1). The first survey was conducted in 1959 by the Ontario Department of Planning and Develop- ment, wherein five sites on the Lynde Creek and one site on Pringle Creek produced Redside Dace (see Table 1 site codes preceded by “OPD”). A second survey conducted by Tumey in 1983 noted 10 Redside Dace capture locations on the Lynde (see Table .1site codes. preceded by: “CLOCA”). The third survey produced one confirmed (Royal Ontario Museum) Lynde Creek specimen taken by Niblett Environmental Associates in 1997 (see Table 1 site code preceded by “NEA”). The fourth, and most recent study conducted by EcoTec in 1999 noted Redside Dace within the Lynde Creek at two sites along the Highway #7 corridor (see Table 1, site codes preceded by “EcoTec’”’). Holm and Crossman (1985) resurveyed three of the 1959 survey loca- tions, and one new site, but no Redside Dace were found. They suggested that in Lynde and Pringle creeks; (1) there has been a significant decline of C. elongatus in the waters sampled, (2) C. elongatus has been seriously limited or extirpated from Pringle Creek, (3) either the number of individuals, or the area of the watershed occupied by the species, has declined in Lynde Creek, and (4) there appears to be healthy populations in other watercourses. The rec- ommendations from this survey concluded; “Urbanization definitely seems to reduce the area of any watershed which remains suitable to C. elonga- tus. It is recommended for streams with known pop- ulations of C. elongatus that wherever possible, the stream-side vegetation be maintained and instream cover and obstructions such as logs not be removed (i.e., channelization be avoided), this will maintain the pool-riffle habitat and supply a source of terres- trial insects — an important food source for the red- side dace”. According to these conclusions and recommen- dations, the decline or extirpation of Redside Dace in the Lynde and Pringle creek systems appears imminent. Lynde and Pringle creeks have been sub- ject to many stresses over the years including urban development and subsequent channelizations and stormsewer outfalls. Many other groups have sur- — veyed Lynde and Pringle creeks in the vicinity of many known Redside Dace occurrences (CLOCA 1996-2000) and have not captured any specimens since 1996, with the exception of the Niblett Vol. 116 Environmental Associates survey (1997). Redside Dace occurrences had not been previously reported at the two sites studied by EcoTec (1999*) even though the sites had been surveyed prior to 1999. Contemporary Study Results A total of 20 Redside Dace were collected at four of the 20 historical collection sites (Table 1) and specimens were verified as C. elongatus (ROM Accession #6844). The average total length was 80.5 mm (ranging between 48-101 mm) and the average weight was 4.94¢ (ranging between 2-10.1 g) for all samples. At the sites where Redside Dace were captured, the following consis- tent characteristics of the aquatic and riparian habi- tat were noted; 2"4 or 34 order streams, a relatively steep gradient (1.0—1.59%), riffle pool sequences, cool water (summer temperatures between 16.0°C and 24.0°C), a streamside vegetation made up of grasses providing 100% canopy cover, a lack of instream woody debris, a surrounding land use of reclaimed agricultural or recreational (golf course) and variable substrata including; silt, sand, cobble and rock. Discussion Existing and Potential Habitat The four capture sites contained similar habitat. It was determined that this habitat type, with some exceptions (see below), could be identified in a desktop exercise using rectified colour infrared aerial photography (Ontario Ministry of Natural Resources 1997). Reclaimed agricultural areas that were not excessively forested and had riparian veg- etation consisting of mainly grass species were denoted as potential habitats (Figure 1). The find- ings derived from this exercise do not preclude other areas and habitat types from being Redside Dace terrestrial habitat, rather, they represent areas within Lynde and Pringle creeks that contain a habitat type that has been proven to support Redside Dace. However, there are limitations in the “desktop” method of habitat typing: some areas of identified potential habitat are either within an ephemeral or intermittent watercourse, or do not have sufficient flows to support Redside Dace (Parker et al. 1988); other areas may have either thermal characteristics outside Redside Dace toler- ance range, or substrates and riffle/pool characteris- tics not normally associated with Redside Dace habitat (Scott and Crossman 1973). Furthermore, fragmented areas of potential habitat are evident within some tributaries and it is unknown if these pockets provide enough habitat to be considered “potential”. For the purpose of this study, these areas are included as the potential for Redside Dace habitat is evident. It would appear that Redside Dace populations are becoming more restricted within Lynde Creek and are either severely limited er | 2002 ANDERSEN: REDSIDE DACE IN LYNDE AND PRINGLE CREEK WATERSHEDS 79 TABLE 1. Fish species captured during Redside Dace sampling conducted in Lynde and Pringle creeks in 1999 and 2000. oO as) ‘= > = LY @) _ < ee Ga pee vA beech een By Soe law Boe. oS rR ety a ae = rene RC Be ie Se relatey "RS. oy tee eas ~Q ar ei BS oe vo ten aS eee Cg = Vette Eel pet & Fe a” AS ay eee ; 2m Se) La, SR: Sa & ee &) Species Salmonidae Oncorhynchus mykiss 18 4 agar | Salvelinus fontinalis 1 Catostomidae Catostomus commersoni 9 MO ty 2 an Die 4 Cyprinidae Phoxinus eos 1 7 1 Clinostomus elongatus Luxilus cornutus 1 Symes | Pimephales promelas 11 4 Rhinicthys atratulus P26" Sl "64 58° he 23) 81 Rhinicthys cataractae 4 8 mien) 2 Semotilus atromaculatus 17 85 635.) Pio. cee 37 Gasterosteidae Culaea inconstans 3 1 2 Centrarchidae Micropterus salmoides 1 Lepomis gibbosus Z 1 Percidae Etheostoma caeruleum Ss “iy 2 Etheostoma nigrum er 14°22 Ss 20 8 Cottidae Cottus bairdi 4 16? "4 1 Total Number of Specimens 162 145 193 167 268 82 143 Total Number of Taxa 6 8 dB, 18, 9. 9 or absent within Pringle Creek. Although the range of occupation of the Redside Dace within Lynde Creek has been reduced, those populations residing in suitable habitat appear to be healthy based on specimen condition and numbers captured (Table 1). According to historical field notes, the land use surrounding each capture site has remained rela- tively unchanged since first being sampled. Those sites that have seen a significant land use change (i.e., from forested to agricultural or vice-versa) since the time of the first sample event did not con- tain Redside Dace. It would appear that any changes to the above noted habitat characteristics would result in a absence of Redside Dace capture. One example of this was discovered north of the Hamlet of Brooklin wherein a capture site Site Code - ~ SB@sessaasae = = S < < hs a f bs as | oO eS gS < r= >A = = Ss ‘= = = = 5 Pp >» ce Fea Aa es ae J =] er — — —_ — — — — Bests wrasaf — N — . «(CO = a. fae © oO Oo oO & oO VU VU UO & = => © OB Om 0 OO Oe 8s oO oO A. ee pe ae ee iS ea] mo O @w UO we VO UO U UO @ = 4 A) sG60-0 2 55 1 4 1 7 , 68 7 2 95 20 - 1 34 13 3 1 2295 5 = 20 r 3 2 = 48 oF 631 66 SE 32 =e 29 (24 22 «25 ee 791 D) 1 21 54 Pe 52 42 20 me gs 5 1 60 659 6 1 3 3 11 20 2 16 2 3 177 1 a) a aie 77 134 136 78 69 65 121 50 75 30 52 95 2065 . 7 6 Me 4m 67 4 «5 me 16 described as agricultural in 1959 has been reclaimed by an Eastern White Cedar forest by 1999 and did not display Redside Dace capture. Development in and around the Hamlet of Brooklin has limited the range of the Redside Dace within this area, possibly due to increased sediment load- ing and turbidity, riparian vegetation removal and temperature changes. Population Management Considerations Populations of Redside Dace throughout their Ontario Range could be subject to a similar exercise as reported here to determine the Provincial popula- tion status in a consistent manner. Redside Dace populations would benefit throughout their range if the strictest levels of sediment and turbidity controls are utilized during residential and agricultural devel- 80 THE CANADIAN FIELD-NATURALIST opment. Further areas of study should give consider- ation to population estimation, measurements of; physical habitat, thermal condition nutrient status, dissolved oxygen and turbidity, and the relationship of migration barriers to Redside Dace populations. Acknowledgments It is with gratitude I thank everyone who endeav- ored to help me “look for little red fish”; Ian Kelsey, Owen Kelsey, Mike Wilson, Cyndee Pettifer, Todd Langley, Jay Howleg, Erling Holm, Jill McColl, Lori Manzon and the Central Lake Ontario Conservation Authority. Documents Cited [marked * in text] Archives of Ontario. 1959. Ontario Department of Plan- ning and Development Stream Survey Forms, RG1, HB Boxes 2-3. Toronto, Ontario. CLOCA (Central Lake Ontario Conservation Author- ity). 1996-2000. Stream habitat assessment summaries for the CLOCA Watershed. Central Lake Ontario Conservation Authority, Oshawa, Ontario. EcoTec Environmental Consultants. 1999. Fisheries Inventory and Assessment Agreement #2005-A-000017. Ministry of Transport, Toronto, Ontario. Holm, E., and E. J. Crossman. 1986. A report on a 1985 attempt to resurvey some areas within the Ontario distri- bution of Clinostomus elongatus, the Redside Dace and to summarize previous records. Unpublished. Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Ontario. Tumey, P. R. 1984. An inventory and assessment of four streams within the Central Lake Ontario Conservation Authority. Unpublished. Central Lake Ontario Conser- vation Authority, Oshawa, Ontario. Vol. 116 - Literature Cited COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2000. Canadian species at risk, yearly report. Canadian Wildlife Service, Environment Canada, Ottawa, Ontario. COSSARO (Committee on the Status of Species at Risk in Ontario). 2000. Index list of vulnerable, threatened, endangered, extirpated or extinct species of Ontario. Queen’s Printer for Ontario, Toronto, Ontario. Daniels, R. A., and S. J. Wisniewski. 1994. Feeding ecol- ogy of Redside Dace Clinostomus elongatus. Ecology of Freshwater Fish 1994 3:176-183. Novinger, D. C., and Thomas G. Coon. 1998. Behavior and physiology of the Redside Dace, Clinostomus elon- gatus, a Threatened Species in Michigan. Environmental Biology of Fishes. Netherlands 57: 315—326 2000. Page, L. M., and B. M. Burr. 1991. A field guide to freshwater fishes. Peterson Field Guide Series, Audubon Society. Houghton Mifflin Company, New York, New York. 432 pages. Parker, B. J., P. McKee, and R. R. Campbell. 1988. Status of the Redside Dace, Clinostomus elongatus, in Canada. Canadian Field Naturalist 102: 163-169. Scott, W. B., and E. J. Crossman. 1973. Freshwater fish- es of Canada. Bulletin 184, Fisheries Research Board of Canada, Ottawa, Ontario. 966 pages. Stanfield, L., M. Jones, M. Stoneman, B. Kilgor, J. Parish, and G. Wichert. 1997. Stream assessment pro- tocol for Ontario. Great Lakes Salmonid Unit, Ministry of Natural Resources, Picton, Ontario. 155 pages. Trautman, M. C. 1981. The fishes of Ohio with illustrat- ed keys. Ohio State University Press, Columbus, Ohio, 683 pages. Received 19 February 2001 Accepted 22 January 2002 Aquatic Leaves and Regeneration of Last Year’s Straw in the Arctic Grass, Arctophila fulva SusAN G. AIKEN! and ROSEMARY A. BUCK? 1Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada; saiken@mus-nature.ca 2156 Dalton Trail, Whitehorse, Yukon Y1A 3G2 Canada Aiken, Susan G., and Rosemary A. Buck. 2002. Aquatic leaves and regeneration of last year’s straw in the arctic grass. Arctophila fulva. Canadian Field-Naturalist 116(1): 81-86. Aquatic leaves are reported and described for the grass Arctophila fulva. Re-growth from apparently dead, previous sea- son’s straw that had over wintered in the Arctic, is reported. Observations on transplanted material and field observations documented that new shoots grew from the apex, and nodes of previous season’s stems. Strong underground rhizomes, and roots developed resulting in propagules able to develop into new plants. The previous season’s straw that regenerates is probably detached by animals mid-season, before food reserves in the stems are withdrawn to the rhizomes and roots. Such re-generation of detached straw that has over wintered has not been reported previously for any other grass species. Key Words: Poaceae, Arctophila fulva, aquatic leaves, vegetative proliferation, Banks Island. The grass Arctophila fulva (Trin.) Rupr. has a low Arctic circumpolar distribution (Porsild 1957; Tzvelev 1976; Aiken et al. 1996, 2000*). In North America, it occurs in Alaska, the Yukon, the Northwest Territories, Nunavut, Ontario, Quebec, and Labrador. The northern-most record for the species in the Western Canadian Arctic Archipelago is from Prince Patrick Island, 76°12’N, 119°25’W, (Canadian National Herbarium (CAN) 220464). Plants grow in shallow, standing water or in wet marshy areas where they often form more or less pure stands generally 20-50 (and occasionally up to 100) cm high. The species is easily recognized because, unlike any other grasses or sedges that occur in the same areas, the mature plants have leaves that are conspicuously larger towards the top of the plant than those at the base (Aiken et al. 2000*). The occurrence of aquatic leaves in grasses is so rare that this is not one of the 1087 characters con- sidered for the 9593 taxa in World Grass Species (Clayton 1999*). In Canada, wild rice taxa in the genus Zizania L. and Signal Grass Pleuropogon sabinei R. Br. were the only grasses found to devel- Op aquatic leaves in the survey done for Watson et al. (1986). Porsild (1957) noted that A. fulva “near its north- ern limit is often sterile, propagating itself vegeta- tively”. He gave no information on the nature of the proliferation and did not describe the propagules. Vegetative proliferation by bulbils, formed in place of flowers, occurs in the North American arctic grasses Poa pratensis (L.) subsp. alpigena (Fr. ex Blytt) Hiitonen and Festuca viviparoidea Krajina ex Pavlick s.]. (Aiken et al. 2000*). Vegetative prolifer- ation by fragmentation of stolons and vertical shoots, as a result of goose foraging, appears to be mainly responsible for the spread of Puccinellia phrygan- 81 odes (Trin.) Scribn. and Merr., a grass not known to set seed, and the sedge Carex subspathacea Wormsk. that rarely sets seed (Chou et al. 1992; per- sonal communication J. Cayouette 2002). In these species the establishment of the propag- ules occurs in the same growing season. Aerial stem fragments can be used as cuttings to produce new plants of Bamboos (sample bought in Ottawa February 2002), but this is not known for any native grasses that occur in Canada, and is unlikely as the stems are annual. In July 1999, Arctophila fulva was common in Aulavik National Park, Banks Island, N.W.T., Canada along the Thomsen River, between 913° 13.49" Ny 199°32i12) Wy andi 75°36" N, 120°2’ W, as well as at other sites in the park. Methods At the site on Banks Island, beside the Thomsen River (73°46’ N, 119°56.7’ W), on 10 July 1999, plants were growing in a series of shallow pools 10-70 cm deep surrounded by wet sedge meadows predominantly of Eriophorum angustifolium Honck. In this pool, most of the new season’s growth was submerged leaves that if they reached the surface, floated parallel to it. Some plants had culms emerg- ing from the water and developing short, aerial leaves. A sample of each growth form was collected and preserved as a herbarium specimen that was deposited at the Canadian National Herbarium (CAN 582410, Figure 1). Beside the tundra pools were pieces of detached straw of the previous season’s growth that were often lying on the damp moss, Hamatocaulis verni- cosus (Mitt.) Hidenas. Most of the straw was dead. Occasionally, new season’s growth, in the form of leafy shoots, was observed at the top of some of the 82 THE CANADIAN FIELD-NATURALIST Vol. 116 FiGURE 1. Herbarium specimen of plants collected N.W.T., Banks Island, Aulavik National Park, 73°46’ N; 119°56’ W, 10 July 1999, S. G. Aiken 99-230 (CAN 582410). Left, submerged plant with thin, flexuous, aquatic leaves and long rhizomes; right, plant devel- oping characteristically distichous, more rigid, emergent leaves. pieces of stranded straw. Samples were collected and preserved as a herbarium specimen (CAN 582408, Figure 2). Twenty-six additional detached stems with new growth, along with some of the moss on which stems were lying, were collected. They were transported to Banks Island, Sachs Harbour, 71°59'N, 125°20’ W, on 13 July 1999 where the straw was sorted into 13 approximately matched pairs of stems of similar length, width, and shoot development. One set remained in Sachs Harbour and the other set was transported to a private home in Ottawa-Carleton, Ontario, 45°16’ N, 75°46’ W. In both locations, the moss was placed in the bottom of a shallow container and the stems were laid on top. Pond or rainwater was used to saturate the moss to duplicate the conditions where the samples were col- lected. The containers were placed outdoors in Sachs Harbour on 14 July, where temperatures were 1—3°C, and there was 24-hour daylight and in Ottawa on 16 July, where temperatures were 17-32°C, and there was a day-night cycle. Observations were made as the season progressed. When it appeared that lack of nutrients may be limit- ing growth, a 10 cm layer of soil was placed under the moss on which the culms were lying. Results Description of aquatic leaves in A. fulva. Aquatic or submerged leaves are long, narrow, and pale pinkish brown. They look similar to the aquatic leaves of the grass Pleuropogon sabinei FIGURE 2. Voucher herbarium specimens collected N.W.T., Banks Island, Aulavik National Park, 73°46’ N; 119°56’ W, 10 July 1999, S. G. Aiken 99-230c (CAN 582408). Left, two typical pieces of previous season’s straw 25-30 cm long. Right, a stem approximately 1 m long, developing new season’s growth at the apex. Note that the remains of the previous season’s leaves approximately 15 cm long have ends that might be interpreted as chewed. R.Br. (image, Aiken et al. 2000*). The leaves were suspended in water like those of the aquatic leaves of wild rice Zizania palustris (Aiken et al. 1988). Submerged leaves have sheaths that are translucent, pale pinkish brown, closed to near the apex. Cells are developed on 50-60% of the sheath circumfer- ence. These are larger in the middle of the tissue and small before a thin translucent membrane that forms the remaining portion of the circumference. In sur- face view cells are readily visible, 24.5 mm long, 0.2—0.3 mm wide, and with prominent end walls. Ligules are adnate to the blade for 3-6 mm and taper to an acute tip from the blade margins to the mid- vein. The free ligule surface is uniformly about 0.5 mm wide, with the abaxial surface glabrous and the margin entire. The ligules are more like those of the Cyperaceae than the Poaceae. Collar position is inconspicuous. Blades are 8-25 cm long, 0.1—-0.4 cm wide, translucent, pinkish brown and limp if removed from the water. Leaf cells, visible in sur- face view, are up to 7 mm long, 0.2-0.3 mm wide with prominent end walls. In leaf cross section, the midvein is slightly larger than the 8-14 adaxial to abaxial girders, 4-7 on either side of the midvein. The adaxial surface epidermis is one cell thick; the abaxial surface has a single layer of colorless epider- mal cells and above them a single layer of cells con- taining chlorophyll. Between the epidermi and the girders are large air spaces. There is a slight develop- 2002 ment of cell thickening at both ends of each girder, but this is not enough to influence the outline of blade surfaces that are uniform in cross section. The aquatic leaves observed when the specimens were collected have not been described previously in North American plants and none was found among circumpolar specimens of A. fulva examined by Aiken at Kew Herbarium (England, October, 2000). Transition and emergent leaves These were beginning to form on shoots that had started to emerge by 10 July 1999 and had relatively stiff leaves less than 10 cm long above the water sur- face. Leaves formed near the water surface were opaque and greenish brown. Aerial leaves of A. fulva have opaque sheaths that are green, brownish green, or sometimes reddish, and open to the base. The margins are not conspicuously membranous except near the sheath apex where the ligule margins are decurrent; cells and cell end walls are not visible. Ligules are free from the surface of the blade, 2—3 (-4) mm long slightly higher at the blade margins with the abaxial surface minutely scaberulous and the apex truncate, lacerate, and ciliolate. The ligule is similar to that found in some other arctic Poaceae (e.g., Arctagrostis latifolia Aiken et al. 2000*). Collars are conspicuous as a zone of contrasting colour on the abaxial surface between the sheath and the blade. They become more conspicuous in older leaves, up to 3 mm wide at the margins, and narrow at the midvein. Blades of the first aerial leaves are 3.54 (-6) cm long and become 10—15 cm long near the inflorescence, 0.2—0.8 cm wide, opaque, green, spreading stiffly at angles of 45—65° from the culm. The position of the vascular bundles is conspicuous as ribs on both leaf surfaces, with a lighter colour over the ribs, but leaf cells are not visible. In leaf cross section, the midvein is often inconspicuous in surface view, only slightly larger than the 6-23 adaxial to abaxial colorless girders. The leaves have typical C, anatomy with zones of chlorenchyma between the bundles. Vascular bundles in girders are slightly deeper than the chlorenchyma zones between the bundles. On top and bottom surfaces of the girders are prominent, bubbly zones of compact colorless parenchyma. In these positions, the outline of the leaf cross-section has conspicuous contours on both surfaces and is 3—4 times as wide as in the zones between ribs. Last year’s dead straw Detached stems that had blown to the shoreline of the tundra were usually 20-50 cm long, but one mea- sured 1 m long (CAN 582408, Figure 2). This stem was conspicuously longer than any other grass stems seen in Aulavik National Park and may have been growing in relatively deep water. There was no evi- dence that the A. fulva stems had flowered the previ- ous season. Rather, where flowering may have AIKEN AND BUCK: AQUATIC LEAVES AND REGENERATION 83 occurred the remains of the previous season’s leaves might be interpreted as having been chewed (Figure 2). A photograph of the new growth developing at the end of the detached stems shows the colour con- trast between the previous season’s straw colored remains and the news season’s growth that is both reddish and fresh green (image Aiken et al. 2000*). On 10 July, none of the nearly 30 sprouting stems collected had developed roots. Ontario, Ottawa-Carleton. Maximum temperatures between 16—23 July 1999, were between 27—32°C. By 18 July, one transplanted stem had sprouted a new shoot from a stem internode, and two days later, the first root appeared. Five days later, two of the termi- nal growing shoots were more than 10 cm high with new green growth. By 23 July, there were 24 actively growing shoots, the original terminal 13 shoots and 11 new shoots developing at culm nodes. Some were less than 1 cm long; the two longest shoots were about 15 cm long and curved upwards (Figure 3A). After this, no increase in height occurred in spite of warm temperatures in August. At the end of August the largest shoots were collected and preserved as herbarium specimens. Significant development of rhizomes and roots occurred below the moss surface during August (Figure 3B) and there was an increase in the number of new shoots produced at nodes along the previous season’s growth until 30 August when a total of 51 shoots were present. Examination of a lon- gitudinal section through apparently dead straw of the previous season’s growth (Figure 4), indicated the stems were brown outside and inside, with little culm wall, and hollow on the inside except at the narrow node from which the new shoots were developing. On 30 August, the largest new stem that had devel- oped was harvested and preserved as CAN 582407. Banks Island, Sachs Harbour. Maximum tempera- tures between 14-24 July 1999, were between 2.6-13°C. By 24 July, many shoots had begun to grow and had red and green leaves. By 24 July, the longest shoot and root were 5 cm and 7 cm long, respectively. There followed two weeks of sunny weather with no rain during which the water in the container dried out, the moss turned brown and many shoots died. By 11 August, there were still 13 shoots alive, some terminal and some developing at nodes. The longest shoot was 8 cm and that was the maximum height reached. By 23 August, there was little sign of more above ground growth and on 31 August, the longest shoot was harvested and pre- served as CAN 582556. Additional field and herbarium observations Observations and voucher specimens were col- lected in 1999 and 2000 in the course of Parks Canada patrols made in Aulavik National Park. Figure 5 was photographed on Banks Island, at Castel Bay, 74°12'N, 119°37'W, on 1 August 1999. 84 THE CANADIAN FIELD-NATURALIST . Vol. 116 FiGuRE 3. A. Close-up of the terminal end of a leafy shoot approximately 15 cm high that had developed in Ontario by 1 August 1999, from a vegetatively propagating piece of straw collected on Banks Island. Scale bar in cm. B. Close-up of new season’s underground growth developing from a node of a previous season’s culm (ps). Note branching rhizome (rz) with a ring of roots (r) developing from a node (n). It shows a vegetative propagule that had developed vigorously growing rhizomes and roots. There are vouchers for observations made at other three other locations in Auluvak National Park, CAN numbers 582894 — 582896. A table summarizing other field observations that were made during Parks Canada patrols in 1999 and 2000 has been place with the specimens in the A. fulva folder, along with notes doc- umenting the occurrence of Muskoxen (Ovibos moschatus Zimmerman) and Brant Geese (Branta bernicula L.) in the area. In 2001, a field trip in July and early August that focused on collecting A. fulva and Duponita fisheri R.Br., visited Churchill, Manitoba; Coral Habour, Southampton and Cornwallis Islands, Nunavut; Tuktoyaktuk, and Prince Patrick Island, Northwest Territories. At none of these places was there any evidence of aquatic leaves or vegetative propagules in either grass species. Discussion Aquatic leaves The aquatic leaves in A. fulva observed on Banks Island were possibly more obvious because plants were growing in water that was relatively deep, not much above freezing, and was still cold around 10 July. Initial growth had been slow and in water where the level had not dropped through drying out as occurs in shallow ponds on Continental North America. The first author observed A. fulva in a drained lake-bed on Richard’s Island, Northwest Territories, 69°20'’N, 134°30’'W, in 1988, where it was the dominant species. The plants were growing in damp moss and mud, where there had probably not been much depth of water in the spring, tempera- tures had been as much as 15°C warmer on average than on Banks Island during June, and growth of aerial stems appeared to have been rapid. If the rela- tively delicate submerged leaves had ever developed, they were no longer visible. Propagules That reserves in detached stems were utilized to develop new shoots from living cells that had not been killed in the severe winter is a phenomenon not known in other Arctic grasses. The vegetative devel- opment of the transplanted stems kept on Banks Island and those in Ottawa-Carleton, as well as the vegetatively propagating specimen found at Castel Bay (Figure 5) and other field observations indicate that under some environmental conditions A. fulva can propagate vegetatively from detached stems. 2002 FiGuRE 4. A specimen cut longitudinally through the node where a new shoot (ns) was developing. Upper por- tion with the base of the new leafy stems (ns) was arising at the node (n) of an otherwise hollow stem. Lower portion is the other side of the stem above. The previous season’s stem (ps) is hollow, and the node (n) is a solid partition across the hollow stem. The lack of roots on the sprouting shoots when they were collected was probably a reflection of an environment where moisture was not limiting growth. Early in development, the need of the straw of A. fulva to develop photosynthetic tissue was apparently greater than the need to develop roots. Although a very different system, the seeds of wild rice (Zizania palustris) germinating where water is not limiting, also develop photosynthetic tissue before producing roots. (Aiken, 1986). The contrast in the growth response between the samples observed on Banks Island and those observed in Ottawa probably reflects the difference in tempera- tures at the two locations. The difference in light quality or day-length may have been responsible for the new growth in Ottawa quickly turning green, while the new growth in plants on Banks Island remained reddish. In the Ontario transplant, A. fulva behaved as a periodic species (that is, one in which development is halted at a particular stage even when abundant opportunity for further growth remains (Sorenson 1941). It might have been tempt- ing to interpret the plants as a genetically dwarfed arctic race of the species towards the northern limit AIKEN AND BUCK: AQUATIC LEAVES AND REGENERATION 85 FIGURE 5. A piece of previous season’s straw successfully vegetatively proliferating. It had two rhizomes up to 12 cm long by 1 August, on Banks Island on the coast of Castel Bay, 74°12’ N; 119°37' W. of its distribution (Saville 1972) had it not been for the previous season’s stem 1 m long (Figure 2). The detached stems behaved somewhat like detached stems of Eurasian Water Milfoil (Myriophyllum spicatum L.) that will sometimes strand on damp mud, put out roots, and develop small terrestrial plants. But in that species, the growth occurs in the same growing season (Aiken et al. 1979). The specialized over-wintering buds (turi- ons) of the native water milfoils (Myriophyllum sibiricum Komorov and M. verticillaturm L.), propa- gate similarly (Aiken and Waltz 1979; Weber and Nooden 1974), but after food reserves have been stored in them as they develop. Towards northern limits of distribution Porsild’s (1957) idea that plants of A. fulva grow- ing near the northern limit of distribution of the species that had not flowered, may propagate vegeta- tively is supported by herbarium vouchers: (a) CAN 127454 (a collection from Banks Island, 73°24'N, 117° W, A.E. Porsild, 17645, on the label of which he noted that the plants were growing in water by a shallow pond where perhaps they are always sterile) and (b) CAN 127453 (from Victoria Island, near 70°39’ N, 117°24’ W, made on 8 August 1946, A.E. Porsild 17237, on the label of which is noted that the sterile specimens were growing in 18” (45 cm) of water, forming a margin around a pond and that the species is apparently always sterile here). Porsild did not entertain the idea that the flowering inflores- 86 THE CANADIAN FIELD-NATURALIST . cences may have been grazed off and given that the culm leaves become longer near the inflorescence the top of the culm is not as apparent as in other grasses. There are records of A. fulva plants collected at similar latitudes that have flowers CAN 5357995 (Banks Island, Egg River 72°27’ N, 124°36’), CAN 535769 (Banks Island, Shoron Lake, 73°N, 124°18’ W) and CAN 203530 (the northernmost specimen from Prince Patrick Island 76°12'N). Animal grazing Arctophila fulva is utilized as a forage by Muskoxen and Brant Geese (CAN 220464 and Park Canada patrol observations in Aulavik National Park). It is possible that plants had begun to flower, or were flowering, but. the inflorescences were removed by some of these animals, grazing. Gray (1987) described Muskoxen playing in water on Bathurst Isiand (where A. fulva does not occur). He had observed that animals would walk into the mid- dle of a shallow pond, stand in the water and sudden- ly begin to head-toss, whirl around, jump and splash. If A. fulva was growing in such a pond, plants would likely be uprooted by such activities. The tundra pools observed on Banks Island would be ideal for such play and the large Muskoxen population on the island may well explain the large number of detached stems found around the margins of ponds. These may float to the surface, and later be blown to the shoreline of the ponds, where they over-winter. Possibly some stems are detached when they are actively growing and contain food reserves that oth- erwise, later in the season, would have been with- drawn to the rhizomes and roots with the onset of winter. Such reserves appear to have been available to initiate new growth the following season, in some detached stems. On both Banks and Victoria islands, Muskoxen and Brant Geese are common and their activities may contribute to A. fulva occasionally proliferating vegetatively from detached straw. When the phenomenon of straw from one season being able to sprout new plants in the next season was discussed with grass taxonomists at Kew, they were unaware of any other such occurrence and thought it most unlikely to happen in tropical or tem- perate areas (W. D. Clayton, S. M. Phillips, and S. A. Renoize, personal communication March 2002). Acknowledgments The authors acknowledge with thanks the logistic support of this project by Parks Canada, and particu- larly by N. A. Lawrence, the Chief Warden of Aulavik National Park, 1999-2000, Polar Continental Shelf Project, and the Canadian Museum of Nature. They also acknowledge permission given to carry out plant research in the Park by Parks Canada, the Hunters and Trappers Committee, Sachs Harbour, and the Environmental Impact Screening Committee, Inuvik. The research was undertaken on the science license #13055N from the Aurora Vol. 116 Research Institute. For help with producing figures, thanks go to R. Boles and J. Madill. Anonymous reviewers and W. D. Clayton, S. M. Phillips, and S. A. Renoize are thanked for helpful discussions on earlier versions of the manuscript. Documents Cited (Marked * in text) Aiken, S. G., L. L.Consaul, and M. J. Dallwitz. 2000 onwards. Grasses of the Canadian Arctic Archipelago: Descriptions, Illustrations, Identification and Information Retrieval. URL [RTF bookmark start: _H]lt492185404] [RTF bookmark end: _HIt492185404]/delta/arcticf. Hard copy deposited at the Canadian Museum of Nature Library. Clayton, W. D. 1999. World Grasses Database v. 5.1 http://www.rbgkew.org.uk/data/grasses, Literature Cited Aiken, S. G. 1986. The distinct morphology and germi- nation of the grains of two species of wild rice (Zizania, Poaceae). Canadian Field-Naturalist 100: 237-240. Aiken, S. G., L. L. Consaul, and M. J. Dallwitz. 1996. Grasses of the Canadian Arctic Archipelago: a DELTA database for interactive identification and illustrated information retrieval. Canadian Journal of Botany 74: 1812-1825. Aiken, S. G., P. F. Lee, D. Punter, and J. M. Stewart. 1988. Wild rice in Canada. NC Press Ltd. Toronto. 130 pages. Aiken, S. G., P. R. Newroth, and I. Wile. 1979. Biology of Canadian Weeds. 34. Myriophyllum spicatum L. Canadian Journal of Plant Science 59: 201-215. Aiken, S. G., and K. F. Waltz. 1979. Turions of Myriophyllum exalbescens Fernald. Aquatic Botany 6: 357-363. Chou, R., C. Vardy, and R. L. Jefferies. 1992. Establish- ment from leaves and other plant fragments produced by the foraging activities of geese. Functional Ecology 6: 297-301. Gray, D. R. 1987. The Muskoxen of Polar Bear Pass. Co- published by the National Museum of Natural Sciences, National Museums of Canada, Ottawa, and Fitzhenry & Whiteside, Markham, Ontario 191 pages. Porsild, A. E. 1957. Illustrated flora of the Canadian Arctic Archipelago. National Museum of Canada Bulletin Number 146. Saville, D. B.O. 1972. Arctic adaptations in plants. Re- search Branch, Canada Department of Agriculture, Monograph 6. Sgrensen, T. 1941. Temperature relationships and phe- nology of the Northeast Greenland flowering plants. Meddelelser om Grgnland 125: 1-305. Tzvelev, N. N. 1976. Grasses of the Soviet Union [Zlaki SSSR. In Russian.] Nauka, Leningrad. Watson, L., S. G. Aiken, M. J. Dallwitz, L. P. Lefkovitch and M. Dubé. 1986. Canadian grass genera: keys and descriptions in English and French from an automated data bank. Canadian Journal of Botany 64: 53-70 + 2 microfiche Data Banks approx. 900 pages. ‘Weber, J. A. and L. D. Nooden. 1974. Turion formation and germination in Myriophyllum verticillatum; phenolo- gy and its interpretation. Michigan Botanist 13: 151-158. Received 20 February 2001 Accepted 14 March 2002 Records of Northern Mockingbird, Mimus polyglottos, Occurrences in North Dakota During the Twentieth Century LAWRENCE D. Ici! and RON E. MARTIN? ‘Northern Prairie Wildlife Research Center, U.S. Geological Survey, 8711 37th Street SE, Jamestown, North Dakota 58401 USA 716900 125th Street SE, Sawyer, North Dakota 58781 USA Igl, Lawrence D., and Ron E. Martin. 2002. Records of Northern Mockingbird, Mimus polyglottos, occurrences in North Dakota during the twentieth century. Canadian Field-Naturalist 116(1): 87-97. The Northern Mockingbird (Mimus polyglottos) is a common bird in the southern United States that has been expanding its breeding range into the northern United States and southern Canada. During the twentieth century, there were 128 reports of Northern Mockingbird occurrences in North Dakota, including 106 reports during the breeding season (15 April to 31 August) and 22 during the nonbreeding season (1 September to 14 April). The species has been largely absent from North Dakota from January through mid-April. Prior to the 1930s, there was only one record (1916) of the Northern Mockingbird in the state. Observations of Northern Mockingbirds in North Dakota increased markedly between the 1930s and 1990s. On average, there were 0.3 reports of mockingbirds per year in 1931-1940, 0.6 in 1941-1950, 1.1 in 1951-1960, 1.6 in 1961- 1970, 2.4 in 1971-1980, 2.3 in 1981-1990, and 4.5 in 1991-2000. The species has been observed in North Dakota nearly annually since 1958. At least six reports during the twentieth century included evidence of nesting (nests or dependent young). Based on mockingbird records during the twentieth century, we designate the current status of the Northern Mockingbird in North Dakota as a rare spring migrant, rare summer visitant, casual nester, and a casual fall and winter visitant. Key Words: Northern Mockingbird, Mimus polyglottos, distribution and abundance, North Dakota, population status, twentieth century. The Northern Mockingbird (Mimus polyglottos) is | reported occurrences of Northern Mockingbirds in a geographically widespread species with southern North Dakota during the twentieth century, review affinities in North America. Densities of the species the species’ current status and distribution within the are highest near the Gulf of Mexico and decline _ state, and discuss factors influencing or limiting the northward and westward (Price et al. 1995). During _ species’ expansion into the state. the twentieth century, the species expanded the northern limits of its range into the northern United Methods States and southern Canada (Derrickson and Records of Northern Mockingbird occurrences in Breitwisch 1992). Although mockingbirds are con- North Dakota were summarized from published and sidered permanent residents throughout much of — unpublished reports, including those in Bird Lore, their historical range, mockingbirds are believed to Audubon Magazine, Audubon Field Notes, American be migratory in the northern portion of their range, Birds, and National Audubon Society Field Notes, including newly colonized northern regions (Brazier and those filed with the North Dakota Birding 1964; Stiles 1982: David et al. 1990; Derrickson and Society (R. E. Martin and G. B. Berkey, North Breitwisch 1992). Dakota Birding Society, unpublished data). We also Stevens (1947, 1948) summarized some of the included data from Christmas Bird Counts (1909- earliest records of Northern Mockingbirds in North — 1999; http://birdsource.cornell.edu/cbc/), North Dakota. Brazier (1964) reviewed the range expan- American Breeding Bird Surveys (1967-1999; Sauer sion of the Northern Mockingbird in the northern et al. 2000: U.S. Geological Survey, Laurel, Great Plains and provided updated and new informa- Maryland, unpublished data), and banding records tion on the Northern Mockingbird in North Dakota. from the Bird Banding Laboratory (1955-1998; U.S. Stewart (1975) summarized historical occurrences of | Geological Survey, Laurel, Maryland, unpublished the species and categorized the Northern data). We defined a record as an occurrence of an Mockingbird as a hypothetical breeder in the state. individual, a breeding pair, or a group of Northern Little has been published on mockingbirds in North Mockingbirds. Records were summarized by date Dakota since Stewart’s book. There has been a sub- (decade and month) and location (county) of the stantial increase in the number of reports of Northern —_gbservation. Details on nests or dependent young Mockingbirds in North Dakota (e.g., Johnson and also were noted. Exact locations or dates are not Johnson 1976), including confirmed nesting records known for a few records. Although some observa- (e.g., Child et al. 1980). In this paper, we summarize tions occurred over multiple days, weeks, or months, 87 88 THE CANADIAN FIELD-NATURALIST | eg Breeding Season 2) = Oo a Nonbreeding Season Ic E ® ” OQ O uw (e) i. ® 2 = = Fl W0t TST ~ 1921," 1931 1941 Decade Vol. 116 isan 1951. 19610-19712 taen FiGuRE 1. Records of Northern Mockingbird occurrences in North Dakota during the twenti- eth century, by decade (e.g., 1901 to 1910, 1991 to 2000). Dark = breeding-season records; light = nonbreeding-season records. summaries within figures represent the date that the individual record was first observed. For purposes of discussion, observations between 15 April and 31 August were considered breeding-season records, and observations between | September and 14 April were considered nonbreeding-season records (Derrickson and Breitwisch 1992) . 60 50 40 30 20 10 Number of Observations a A res ie ae Results During the twentieth century, there were 128 reports of Northern Mockingbirds in North Dakota (Appendix A). The number of individuals detected during a single observation ranged from one to four. Most of the observations involved single birds observed for only one or a few days. So el ee ae Month FIGURE 2. Records of Northern Mockingbird occurrences in North Dakota during the twentieth century, by month. 2002 The earliest confirmed record of a Northern Mockingbird in North Dakota was of an adult bird collected on 23 November 1916 by C. C. Smith on the campus of the University of North Dakota in Grand Forks (Grand Forks County) (Wood 1923). Wood (1923: 77) considered this bird to be an “acci- dental straggler.” To our knowledge, there were no records of Northern Mockingbirds in North Dakota before that date. This observation also represented the first record of the species in the northern Great Plains during the nonbreeding season (Brazier 1964). The second record of a Northern Mockingbird in North Dakota occurred over 17 years later on 17 May 1934 in Valley City (Barnes County) (Appendix A). This observation represented the first breeding-season record in the state and generally marked the beginning of the species’ range expan- sion into North Dakota. Observations of mocking- birds increased markedly between the 1930s and 1990s (Figure 1). On average, there were 0.3 reports of mockingbirds per year in 1931-1940, 0.6 in 1941- 1950, 1.1 in 1951-1960, 1.6 in 1961-1970, 2.4 in 1971-1980, 2.3 in 1981-1990, and 4.5 in 1991-2000. Since 1958, the species has been observed nearly annually in all but 3 of 43 years, with no records for 1962, 1983, and 1989. Only 22 mockingbird occurrences were recorded during the nonbreeding season, mostly between FiGurRE 3. Northern Mockingbird nest found on 25 June 1998 in Kidder County, North Dakota (photograph by L. D. Igl). IGL AND MARTIN: MOCKINGBIRD OCCURRENCES IN NORTH DAKOTA 89 October and December (Figure 2; Appendix A). Mockingbirds were largely absent from the state in January, February, and March. Only two mocking- birds have been observed in North Dakota during those months. Between 30 November 1972 and 14 April 1973, an injured mockingbird was observed in Fargo in Cass County. Between mid-December 1999 and 1 February 2000, a single mockingbird sporadi- cally visited bird feeders about 6.4 km southeast of Jamestown in Stutsman County (M.A. Sovada, U.S. Geological Survey, Jamestown, North Dakota, per- sonal communication). About 83% (106 of 128) of the mockingbird obser- vations occurred during the breeding season (mid- April through August) (Figure 2, Appendix A). Of the 106 reports of mockingbird occurrences in North Dakota during the breeding season, six (6%) involved nests or dependent young. Mrs. R. Nordbye (personal communication in Child et al. 1980) found a nest with young in Parshall (McLean County) in the sum- mer of 1961. Near Richardton (Stark County), J. Hoff found nesting mockingbirds in July 1970, June 1988, and July 1995. Successful fledging of two successive broods in June and July 1980 was reported in Oakes in Dickey County (Childs et al. 1980). L. D. Igl found a breeding pair and a nest containing four eggs in a single-row shelterbelt in Kidder County on 25 June 1998 (Figure 3; Martin 1998). There also are 90 THE CANADIAN FIELD-NATURALIST | Volts 1a BOTTINEAU Lt ROLETTE CAVALIER PEMBINA OWNER 3 ne McHENRY 9 NELSON | GRAND FORKS Id 1 Eps STEELE 1 SHERIDAN] WELLS | FOSTER 2 aaa | TRAILL SD sown Ea 1 LOGAN 1 LAMOURE Th q* | 2 McINTOSH DICKEY SARGENT _——k—_———_= 0 30 FiGurE 4. Records of Northern Mockingbird occurrences in North Dakota during the twentieth century, by county. Asterisk (*) denotes counties where nests or dependent young have been observed. two references to mockingbirds nesting in North Dakota but without specific reference to date: A. C. Fox indicated that mockingbirds have been found nesting in Dickinson (Stark County; Stewart 1975), and R. L. Rytter noted adult mockingbirds feeding young in Kenmare (Ward County). Northern Mockingbirds were observed in 36 (67.9%) of the 53 counties in North Dakota (Figure 4, Appendix A). Although more mockingbird obser- vations occurred in the southern half than the north- ern half of North Dakota (Figure 4), the temporal pattern of expansion into North Dakota was sparse and sporadic rather than progressing from south to north (Appendix A). Counties with larger cities had more records of mockingbirds than less-populated counties. These included Bismarck in Burleigh County, Mandan in Morton County, Fargo in Cass County, Jamestown in Stutsman County, Minot in Ward County, Dickinson in Stark County, and Grand Forks in Grand Forks County. The largest number of records (n = 26) in a county occurred in Cass County, with most occurrences being from Fargo, the largest metropolitan area in North Dakota. The distribution of mockingbird records within the state also may reflect the distribution of observers in North Dakota. For example, there were seven records of mockingbirds in Adams County in south- western North Dakota, all made by C. Griffiths and/or D. Griffiths. The North American Breeding Bird Survey (BBS) is a program of standardized roadside surveys administered by the U.S. Geological Survey and conducted by experienced or knowledgeable volun- teers, usually in June. BBSs have been conducted in North Dakota since 1967; there are currently 44 active routes in the state. Northern Mockingbirds were recorded on 5 of 983 (<1%) surveys of BBS routes that were conducted in North Dakota between 1967 and 1999, including one year each on the Parshall (McLean County), Denbigh (McHenry County), and Sheyenne Lake (Sheridan County) routes and two years on the Bentley (Hettinger County) route (Appendix A). The first occurrence of a mockingbird on a BBS route was on the Parshall route on 19 June 1985. Sponsored by the National Audubon Society, Christmas Bird Counts (CBCs) are conducted annu- ally throughout North America. The main goal of the CBC program is to monitor bird populations during early winter (late December to early January). CBC circles are frequently placed where the greatest con- centrations of wintering birds occur and often are placed near towns and cities. CBCs have been con- ducted in North Dakota since the early part of the twentieth century. The first CBC in North Dakota occurred in Fargo in 1909 (Chapman 1910); one or -more CBCs have been conducted nearly annually in North Dakota since then. Mockingbirds have been reported on 4 of 682 (< 1%) CBCs in North Dakota between 1909 and 1999, including three years on the Bismarck-Mandan CBC and one year on the Fargo- Moorhead CBC (Appendix A). Mockingbirds also 2002 were reported within the Count circle during the week of the CBC but not the day of the CBC in Fargo (Cass County) in 1972 and in Jamestown (Stutsman County) in 1999. The earliest occurrence of the species on a CBC was in Bismarck (Burleigh County) on 20 December 1956. To our knowledge, only four Northern Mockingbirds have been banded in North Dakota. A mockingbird was banded by H. R. Gray in Wilton (McLean County) on 3 June 1943 (Stevens 1948). Of 610 066 birds banded in North Dakota between 1955 and 1998 (all species combined), three were Northern Mockingbirds banded in 1960, 1964, and 1971 by R. T. Gammell and A. M. Gammell (Bird Banding Laboratory, U.S. Geological Survey, Laurel, Maryland, unpublished data). Discussion The available data suggest that North Dakota cur- rently supports a small but growing population of Northern Mockingbirds. Spring arrival in North Dakota begins in mid-April and continues into May (Figure 2), and available evidence suggests that nest- ing occurs largely in June and July. One or two broods are possible, but rare, in North Dakota. Few mockingbirds have overwintered in the state, which, along with the influx of mockingbirds in spring, sug- gests that this species is migratory in North Dakota. However, most of the North Dakota observations were of single birds, and to our knowledge, most birds were observed or present for only one or a few days. Observations of mostly single birds and the general scarcity of nesting suggests that most mock- ingbirds observed in North Dakota were probably wandering rather than nesting birds. Mockingbirds are noted for their behavioral plasticity and wander- ing behavior (Derrickson and Breitwisch 1992). Many records of mockingbird occurrences in North Dakota were from scattered or random obser- vations by one or a few observers. Most observations occurred in spring or summer, and many observa- tions occurred in or near human population centers. Admittedly, these records may be biased by the activity and locations of observers (i.e., most birding occurs in spring and summer and most bird observa- tions are made in residential areas). Apparent increases in North Dakota during the twentieth cen- tury also may reflect higher observer effort or more qualified observers. However, mockingbirds are a conspicuous and easily detected species. Mockingbirds strongly defend their territories during both the nonbreeding and breeding seasons, vocalize loudly and elaborately, and often are associated with human residential areas. Thus, increases in mocking- bird populations based on individual records proba- bly track statewide trends. Changes in mockingbird populations in North Dakota also coincide with similar patterns of range IGL AND MARTIN: MOCKINGBIRD OCCURRENCES IN NORTH DAKOTA 9] expansion in other states and Canadian provinces. During the twentieth century, the species has expanded its range northward in California (Arnold 1980), Michigan (Dziepak 1991), New York (Bull 1974), Ontario (Curry 1987), Quebec (David et al. 1990), Saskatchewan (Smith 1996), and Wisconsin (Robbins 1991). There are scattered records of mockingbirds in states adjacent to North Dakota, including Montana (MBDC 1996), South Dakota (SDOU 1991), and Minnesota (Janssen 1987). In western Illinois, the species was historically an irreg- ular migrant until the mid-1930s, when it became common (Bohlen 1989). Mockingbirds were sparse in Iowa until the 1930s (Spess Jackson et al. 1996); the species is now considered a rare summer resident with higher densities in the southern portion of the state (Kent and Dinsmore 1996). Factors thought to contribute to the species’ north- ward range expansion in North America include increased availability of suitable nesting habitat (trees and shrubs) and increased availability of food (e.g., fruit-bearing trees and shrubs, suet feeders). Northern Mockingbirds favor brushy areas and woodland edges, and often are associated with orna- mental plantings in residential areas (Derrickson and Breitwisch 1992) and shelterbelts and scattered trees and shrubs in agricultural areas (Versaw 1998). The abundance and distribution of woody vegeta- tion have changed dramatically in North Dakota since this region was first settled by Europeans in the mid to late 1800s. Since settlement, there have been repeated efforts to establish tree plantings in North Dakota and other areas in the Great Plains (Hart and Hart 1997). Fire suppression also encouraged the encroachment and establishment of woody vegeta- tion into open grasslands in this region. Between 1935 and 1942, 38 million trees were planted in 13 760 ha of shelterbelts and windbreaks (Haugen 1999). Since the Dust Bowl, more than 142 000 km of field windbreaks and 115 000 ha of protection tree plantings have been established in North Dakota. It is probable that the planting of shelterbelts and wind- breaks in North Dakota in the early part of the twen- tieth century provided the habitat necessary and impetus for the Northern Mockingbird’s northward expansion into North Dakota. Presently, there does not appear to be a shortage of suitable breeding habi- tat for Northern Mockingbirds in North Dakota. Fruit is a critical component of the mockingbird’s diet during the fall, winter, and spring (Derrickson and Breitwisch 1992). Stiles (1982) speculated that the species’ northward expansion into the northeast- ern U.S. may have been related to the plantings of multiflora rose (Rosa multiflora). In the northern Great Plains, mockingbirds have been observed feeding on fruit from several common fruit-bearing trees and shrubs including honeysuckle (Lonicera spp.), Russian Olive (Elaeagnus angustifolia), and 9? THE CANADIAN FIELD-NATURALIST apple (Malus spp.) (Brazier 1964). In other areas, the species has been observed eating fruit from hawthorn (Crataegus spp.), crabapples (Pyrus spp.), and Eastern Redcedar (Juniperus virginiana) (Arnold 1980; Stiles 1982), all of which occur in North Dakota. Dziepak (1991) indicated that suet feeders and ornamental plantings of fleshy-fruited shrubs may have helped the species expand its range into Michigan, but these food resources were not sufficient to sustain the species through Michigan’s severe winters. Northern Mockingbirds are largely absent from North Dakota during the winter. Overwintering also is rare in other northern regions (e.g., Saskatchewan; Smith 1996), suggesting that mockingbirds withdraw from these areas for the winter or succumb to winter conditions. Dziepak (1991) and others have suggest- ed that winter weather is likely the chief factor limit- ing the distribution of this species at the northern edge of its range. Root (1988) proposed that ‘a physi- ological limit on metabolic rate constrains the north- ern limit of wintering North American birds, and demonstrated that mockingbirds typically overwinter in areas that have an average minimum January tem- perature of —7°C or higher. The average January temperature in North Dakota ranges from —16.7°C in northeast North Dakota to —8.3°C in southwest North Dakota (Jensen 1972). Repasky (1991), how- ever, argued that the northern boundary of a species would occur at the point where sufficient food can- not be obtained to offset the greater energy demand of colder temperatures. Brazier (1964) noted many records of dead mockingbirds found during winter in the northern Great Plains. In Illinois, mockingbirds have suffered massive reductions in population dur- ing severe winter weather (Bohlen 1989). The suc- cess of the species’ range expansion into northern areas will likely be determined by the species’ ability to develop stronger migratory habits or to cope with the harsh winter conditions and limited food resources in northern regions. Based on mockingbird records of occurrence in North Dakota during the twentieth century, we des- ignate the current status of the Northern Mock- ingbird in North Dakota as: (1) a rare spring migrant (i.e., a species that occurs annually in North Dakota in low numbers during spring), (2) a rare summer visitant (1.e., a species that wanders or oversummers annually in low numbers in North Dakota during the normal breeding period), and (3) a casual fall and winter visitant (i.e., a species that occurs less than annually in North Dakota during fall or winter). Although there were very few nesting records in North Dakota during the twentieth century, we also recommend that the species’ nesting status in North Dakota be changed from “hypothetical breeder” (Stewart 1975) to “casual nester” (i.e., a species for which a viable clutch of eggs, dependent young in the nest, or dependent young that have left the nest Vol. 116 have been observed less than annually in North Dakota). Continued monitoring of mockingbirds should provide insight into the dynamics of the species’ range expansion into North Dakota and elsewhere in the northern Great Plains. Future increases or changes in the species’ distribution and abundance in North Dakota will necessitate a re- evaluation of the species’ status. Acknowledgments Over the years, many people have contributed records of Northern Mockingbirds in North Dakota. We thank all of them. We are especially grateful to Gordon B. Berkey and David O. Lambeth for their dedication to North Dakota birdlife, for their com- mitment to the North Dakota Birding Society, and for compiling bird records for the state from the mid- 1970s to the mid-1990s. This paper has benefitted from the thoughtful comments of G. B. Berkey, F. R. Cook, A. J. Erskine, C. S. Houston, D. H. Johnson, H. T. Sklebar, and M. A. Sovada. Literature Cited Arbib, R. L., H. Heilbrun, and The Regional Christmas Bird Count Editors of American Birds. Editors. 1973. 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Bird Lore’s tenth Christ- mas Census. Bird Lore 12: 19-36. Child, D., J. Fontaine, T. Gatz, M. Johnson, and J. Oswald. 1980. Northern Mockingbird breeding in North Dakota. Prairie Naturalist 12: 87-88. Cruickshank, A.D. Editor. 1957. Fifty-seventh Christmas Bird Count. Audubon Field Notes 11: 68- 234. » Cruickshank, A. D. Editor. 1964. Sixty-fourth Christmas Bird Count. Audubon Field Notes 18: 77-322. Cruickshank, A. D. Editor. 1970. Seventieth Christmas Bird Count. Audubon Field Notes 24: 101-464. Curry, R. L. 1987. Northern Mockingbird. Pages 334-335 in Atlas of the breeding birds of Ontario. Edited by M. 2002 D. Cadman, P. F. J. Eagles, and F. M. Helleiner. Univer- sity of Waterloo Press, Waterloo, Ontario. David, N., M. Gosselin, and G. Seutin. 1990. Pattern of colonization by the Northern Mockingbird in Quebec. Journal of Field Ornithology 61: 1-8. Derrickson, K. C., and R. Breitswisch. 1992. Northern Mockingbird (Mimus polyglottos). In The birds of North America, Number 7. 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Prairie Naturalist 8: 60. Kent, T. H., and J. J. Dinsmore. 1996. Birds in Iowa. Published by authors, lowa City and Ames, Iowa. 391 pages. Krause, H. 1959. Northern Great Plains region. Audubon Field Notes 13: 41-43. Krause, H. 1960. Northern Great Plains region. Audubon Field Notes 14: 47-52. Lambeth, D. O. 1982. Northern Great Plains. American Birds 36: 190-192. Lambeth, D. O. 1985. Northern Great Plains region. American Birds 39: 69-71. Lambeth, D. O. 1989. Northern Great Plains region. American Birds 43: 121-123. IGL AND MARTIN: MOCKINGBIRD OCCURRENCES IN NORTH DAKOTA 93 Lambeth, D. O. 1990. Northern Great Plains region. American Birds 44: 447-450. Lambeth, D. O. 1991. Northern Great Plains region. American Birds 45: 462-464. LeBaron, G.S. Editor. 1994. The ninety-fourth Christmas Bird Count: December 17, 1993 to January 3, 1994. National Audubon Society Field Notes 48: 355-901. Martin, R. E. 1994. Northern Great Plains region. National Audubon Society Field Notes 48: 311-312. Martin, R. E. 1998. Northern Great Plains region. National Audubon Society Field Notes 52: 470-471. Montana Bird Distribution Committee (MBDC). 1996. P. D. Skaar’s Montana bird distribution, fifth edition. Montana Natural Heritage Program, Special Publication Number 3. 129 pages. Price, J., S. Droege, and A. Price. 1995. The summer atlas of North American birds. Academic Press, New York. 364 pages. Randall, R. N. 2000. Christmas Bird Counts for North Dakota 1999. Prairie Naturalist 32: 115-119. Repasky, R. R. 1991. Temperature and the northern distri- butions of wintering birds. Ecology 72: 2274-2285. Robbins, S. D., Jr. 1991. Wisconsin birdlife. University of Wisconsin Press, Madison. 702 pages. Root, T. L. 1988. Atlas of wintering North American birds: an analysis of Christmas Bird Count data. University of Chicago Press, Chicago, Illinois. 312 pages. Sauer, J. R., J. E. Hines, I. Thomas, J. Fallon, and G. Gough. 2000. The North American Breeding Bird Survey, results and analysis 1966 - 1999. Version 98.1, USGS Patuxent Wildlife Research Center, Laurel, Mary- land. http://www.mbr.nbs.gov/bbs/bbs.html. Serr, E. M. 1976a. Northern Great Plains. American Birds 30: 855-858. Serr, E. M. 1976b. Northern Great Plains. American Birds 30: 969-972. Serr, E. M. 1978. Northern Great Plains. American Birds 32: 1021-1024. Serr, E. M. 1979a. Northern Great Plains. American Birds 33: 782-784. Serr, E. M. 1979b. Northern Great Plains region. American Birds 33: 874-875. Serr, E. M. 1980. Northern Great Plains. American Birds 34: 789-791. Smith, A. R. 1996. Atlas of Saskatchewan birds. Environ- ment Canada-Nature Saskatchewan. Saskatchewan Natural History Society (Nature Saskatchewan), Sas- katchewan, Regina. Manley Callin Series Number 4. Special Publication 22. 456 pages. South Dakota Ornithologists’ Union (SDOU). 1991. The birds of South Dakota. Northern State University Press, Aberdeen, South Dakota. 411 pages. Spess Jackson, L., C. A. Thompson, and J. J. Dinsmore. 1996. The Iowa breeding bird atlas. University of lowa Press, Iowa City. 484 pages. Stevens, O. A. 1947. Mockingbirds in North Dakota. Flicker 19: 19-20. Stevens, O. A. 1948. New and unusual North Dakota trapping records. Auk 65: 136-137. Stewart, R. E. 1975. Breeding birds of North Dakota. Tri-College Center for Environmental Studies, Fargo, North Dakota. 295 pages. Stiles, E. W. 1982. Expansions of mockingbird and multi- flora rose in the northeastern United States and Canada. American Birds 36: 358-364. 94 Versaw, A. E. 1998. Northern Mockingbird (Mimus poly- glottos). Pages 400-401 in Colorado breeding bird atlas. Edited by H. E. Kingery. Colorado Bird Atlas Partner- THE CANADIAN FIELD-NATURALIST ship and Colorado Division of Wildlife. Wood, N. A. 1923. A preliminary survey of the bird life of North Dakota. Museum of Zoology, University of APPENDIX A. Location, date(s), observer(s), season (nonbreeding or breeding), and number of birds observed for reported occurrences of Northern Mockingbirds in North Dakota during the twentieth century. Data were summarized from published and unpub- lished (R. E. Martin and G. B. Berkey, North Dakota Birding Society) reports. No. l. ps Oo Location Grand Forks, Grand Forks County Valley City, Barnes County . Dickinson, Stark County . Dickinson, Stark County . Fargo, Cass County . Rock Lake, Towner County . Minot, Ward County . Wilton, McLean County . Fargo, Cass County . Minot, Ward County . Mandan, Morton County . Kenmare, Ward County . Bismarck, Burleigh County . Fargo, Cass County . Des Lacs National Wildlife Refuge, Ward County . Southeast of Kenmare, Ward County . Mandan, Morton County . Parshall, McLean County . Des Lacs National Wildlife Refuge, Ward County . Ward County . Kenmare, Ward County . Parshall, McLean County . Mandan, Morton County . Ward County . Bismarck, Burleigh County . Fargo, Cass County . Near Binford, Griggs County Date(s) 23 November 1916 17 May 1934 Summer 1938 May 1939 12-13 May. 1942 31 May 1943 1-2 June 1943 3 June 1943 21-28 July 1946 1-2 June 1948 24-25 May 1952 1-14 June 1952 8 June 1952 13 May 1953 4-7 October 1955 18 May 1956 30 December 1956 Vol. 116 Michigan, Ann Arbor, Michigan. Miscellaneous Publi- cations Number 10. 96 pages. Received 21 March 2001 Accepted 22 March 2002 Observer(s) C. C. Schmidt (Woods 1923) Observer not specified (from the files of the North Dakota Birding Society) R. W. Smith (Stevens 1947) R. W. Smith, F. C. Butcher, and F. G. Butcher (Stevens 1947) Mrs. W. E. Brentzel Ambrosen and Dinkins (Hammond 1943) E. A. Hibbard (Stewart 1975) H. R. Gray (Stevens 1948) O. A. Stevens and H. G. Heggeness (Stevens 1947) S. Saugstad (Stewart 1975) R. N. Randall R. T. Gammell and A. M. Gammell (Gammell and Gammell 1952) R. N. Randall (Baumgartner 1952) J. F. Cassel (Gammell and Gammell 1953) H. S. Huenecke (Gammell and Huenecke 1956) V. Rytter (Gammell 1956) Bismarck-Mandan Christmas Bird Count (Cruickshank 1957) 4-11 November 1958 Dr. and Mrs. R. Nordbye; 23 August 1959 6 May 1960 25-28 May 1960 Summer 1961 29 December 1963 | May 1964 10 May 1964 13 June 1965 June 1966 R. T. Gammell and A. M. Gammell (Krause 1959) A. M. Gammell (Krause 1960) R. T. Gammell R. L. Rytter and D. Rytter (Gammell 1960) Mrs. R. Nordbye (personal communication in Child et al. 1980) Bismarck-Mandan Christmas Bird Count (Cruickshank 1964) R. T. Gammell R. N. Randall Mrs. F. B. Scheel (Hatch 1965) R. E. Stewart, Sr. (Stewart 1975) Season* Nonbreeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Nonbreeding Breeding Nonbreeding Nonbreeding Breeding Breeding Breeeding Breeding Nonbreeding Breeding Breeding Breeding Breeding Number of birds one (collected) one one one one one one one (banded) one one four one (banded) one each at three locations nest with young one one (banded) one one —_— a a ee eee (Continued) 2002 APPENDIX A. (continued) No. 28. ao 30. Sb. 32. a3: 34. 35. 36. Location Hankinson, Richland County (T130N, R50W) North of Fargo, Cass County Bismarck, Burleigh County Fargo, Cass County Rhame, Bowman County Fargo, Cass County Arrowwood National Wildlife Refuge, Stutsman County Des Lacs National Wildlife Refuge, Ward County Fargo, Cass County . North of Richardton, Stark County . Audubon National Wildlife Refuge, McLean County . Montpelier, Stutsman County . Bowman, Bowman County . Kenmare, Ward County . Gackle, Logan County . Jamestown, Stutsman County . Fargo, Cass County . Bismarck, Burleigh County . Cooperstown, Griggs County . Fargo, Cass County . Bismarck, Burleigh County . Kulm, LaMoure County . Jamestown, Stutsman County . Fargo, Cass County . Fargo, Cass County . Sargent County . Hensler, Oliver County . Fargo, Cass County . Fargo, Cass County . Crosby, Divide County . Upper Souris National Wildlife Refuge, Ward County . Northern Prairie Wildlife Research Center, Stutsman County . Grand Forks, Grand Forks County . Oakes, Dickey County Date(s) 12 July 1966 30 April-19 May 1967 20 May 1967 22-23 May 1967 20 June 1967 13 September 1967 21 April 1968 8 May 1968 20 December 1969 July 1970 8-9 May 1971 11, 13-15 May 1971 15 May 1971 3 August 1971 16 May 1972 4 June 1972 12 September 1972; 30 November 1972 to 14 April 1973 28 May 1973 6 July 1973 8 August 1973 26 May 1974 10 May 1975 18 May 1976 25 May 1976 22 June to 11 July 1976 29 June 1977 3 July 1977 3 November 1977 5-30 November 1977 A. Lies 19 May 1978 27 May 1979 5-16 July 1979 6-7 May 1980 8 June to 25 July 1980 IGL AND MARTIN: MOCKINGBIRD OCCURRENCES IN NORTH DAKOTA Observer(s) Season* R. E. Stewart, Sr. Breeding (Stewart 1975) R. Kroodsma Breeding R. N. Randall Breeding Observer not specified Breeding (from the files of the North Dakota Birding Society) R. E. Stewart, Sr. Breeding (Stewart 1975) Observer not specified Nonbreeding (from the files of the North Dakota Birding Society) J. T. Lokemoen and Breeding P. F. Springer A. M. Gammell Breeding Fargo-Moorhead Christmas Nonbreeding Bird Count (Cruickshank 1970) J. Hoff Breeding D. C. McGlaughlin Breeding (Houston 1971) L. C. Haynes (Houston 1971) Breeding J. Oberfoel (Houston 1971) — Breeding R. T. Gammell Breeding G. L. Krapu Breeding M. L. Avery (Stewart 1975) Breeding A. M. Bolin (Houston and Nonbreeding Houston 1973), also reported on Fargo-Moorhead Christmas Bird Count during Count week (Arbib et al. 1973) R. N. Randall (Houston 1973) Breeding D. L. Kubischta Breeding Observer not specified Breeding (from the files of the North Dakota Birding Society) B. Quanrud Breeding Observer not specified Breeding (from the files of the North Dakota Birding Society) D. H. Johnson and Breeding J. E. Johnson (Johnson and Johnson 1976) J. F. Cassel (Serr 1976a) Breeding V. J. Schell (Serr 1976b) Breeding R. Schmidt, J. Herman, Breeding and K. Wilson R. N. Randall Breeding C. Brakke Nonbreeding R. A. Stromstad (Serr 1978) — Breeding I. O. Rostad (Serr 1979a) Breeding C. A. Faanes and other Breeding Center staff (Serr 1979b) J. Van Rybroek (Serr 1980) — Breeding D. Child, J. Fontaine, T. Gatz, Breeding M. Johnson, and J. Oswald (Child et al. 1980) Nonbreeding 95 Number of birds one one one one one one one one one two + young one one + one one one (banded) one one one (injured) one one one one one one one two + two nests (totaling 8 young) (Continued) 96 APPENDIX A. (continued) No. Location 62. Fargo, Cass County 63. Grand Forks Air Force Base, Grand Forks County 64. Mercer County 65. Fargo, Cass County 66. Fargo, Cass County 67. Mandan, Morton County 68. Fargo, Cass County 69. Lake Alice, Ramsey County 70. McLean County 71. Hettinger, Adams County 72. Fargo, Cass County 73. New Town, Mountrail County 74. Sioux County 75. McHenry County 76. Grand Forks, Grand Forks County 77. Eddy County 78. Hettinger, Adams County 79. J. Clark Salyer National Wildlife Refuge, McHenry County 80. Richardton, Stark County 81. Fargo, Cass County 82. Grand Forks, Grand Forks County 83. Hettinger, Adams County 84. Lostwood National Wildlife Refuge, Burke County 85. Fargo, Cass County 86. Mercer County 87. Bucyrus, Adams County 88. Kelly’s Slough National Wildlife Refuge, Grand Forks County 89. Fargo, Cass County 90. Kidder County 91. Jamestown, Stutsman County 92. Bowman County 93. Mirror Pool, Ransom County 94. Grand Forks, Grand Forks County 95. Fried, Stutsman County 96. Fargo, Cass County THE CANADIAN FIELD-NATURALIST. Date(s) 3-18 November 1981 19 May 1982 22 July 1982 11 May 1984 21 June 1984 27 October 1984 17 November 1984 2 May 1985 19 June 1985 28-29 April 1986 22 May 1987 22-23 May 1987 24 May 1987 21 June 1987 Late June 1987 5 August 1987 22 April 1988 13 May and 15 May 1988 June 1988 7 June to 11 July 1988 25 October to 17 November 1988 31 May 1990 9 July 1990 10-14 May 1991 29 May 1991 10 June 199] 23 June 1991 11 May 1992 16 May 1992 18 May 1992 24 May 1992 19 July 1992 28 October to 30 November 1992 4 May 1993 10-19 May 1993 97. South of Bismarck, Burleigh County 15 May 1993 Observer(s) M. A. Bergan and C. A. Spurbeck (Lambeth 1982) J. F. Kelly (Berkey 1982) Faanes (Faanes 1983) Bergan and C.A. M. A. C. A. Spurbeck (Berkey 1984) M.A. Bergan and C. A. Spurbeck J. Swanick and O. Swanick (Lambeth 1985) T. Dahlen (Lambeth 1985) R. W. Schnaderbeck R. E. Martin (Parshall Breeding Bird Survey) D. Griffiths and C. Griffiths L. L. Falk R. A. Satermo and B. C. Houser R. Titus R. L. Rytter (Denbigh Breeding Bird Survey) Observer not specified (from the files of the North Dakota Birding Society) R. Hill D. Griffiths and C. Griffiths (Berkey 1988a) W. Berg, R. E. Martin, and G. B. Berkey (Berkey 1988a) J. Hoff R. Kain (Berkey 1988b) D. O. Lambeth (Lambeth 1989) D. Griffiths and C. Griffiths (Lambeth 1990) R. K. Murphy J. O. Hersch (Lambeth 1991) R. E. Martin (Lambeth 1991) D. Griffiths and C. Griffiths E. E. Freeberg L. L. Falk M. A. Otnes J. T. Lokemoen Sick M. A. Otnes . O. Lambeth J. T. Price and A. Price D. Wiesenborn R. H. O’Connor, G. E. Nielsen, and . A. Otnes Martin and G. B. Berkey Season? Nonbreeding Breeding Breeding Breeding Breeding Nonbreeding Nonbreeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Nonbreeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Nonbreeding Breeding Breeding Breeding Vol. 116 Number of birds one one one one one (injured) one one one one one one one one one two one one two two + nest (with eggs) one one eee ee eee (Continued) 2002 APPENDIX A. (continued) No. 98. 99. 100. 107. . Hettinger, Adams County . Northern Prairie Wildlife 110. tii. 112. ifs, 114. 115. 116. 117. 118. 119. 120. 121. 122. £23. 124. 125. 126. 127. 128. “Observations between 15 April and 31 August were considered breeding-season records, and observations between | September and Location Sheridan County Hettinger County Lostwood National Wildlife Refuge, Burke County . Bismarck, Burleigh County . Hettinger, Adams County . Minot Sewage Lagoons, Ward County . Bottineau County . Grand Forks, Grand Forks County . Theodore Roosevelt National Park (South Unit), Billings County McHenry County Research Center, Stutsman County North of Cayuga, Sargent County Arrowwood National Wildlife Refuge, Stutsman County Near Richardton, Stark County Mandan, Morton County McKenzie County Arrowwood National Wildlife Refuge, Stutsman County Fargo, Cass County Northern Prairie Wildlife Research Center, Stutsman County Fargo, Cass County Richardton, Stark County Mandan, Morton County Grand Forks County Hettinger County Kidder County New Hradec, Dunn County Portland, Traill County Adams County Southeast of Jamestown, Stutsman County Fargo, Cass County Date(s) 13 June 1993 19 June 1993 27 June 1993 20 December 1993 11 May 1994 18 May 1994 22 May 1994 26 May 1994 5 June 1994 5-12 June 1994 13-29 July 1994 7 September to mid-November 1994 10 May 1995 13 May 1995 27 July 1995 15 November 1995 25 November 1995 10 May 1996 24 May 1996 10 June 1996 28 June 1997 19 July 1997 15 April 1998 16 and 18 May 1998 1 July 1998 25 June 1998 27 October 1998 30 October 1998 18 August 1999 Mid-December 1999 to 1 February 2000 9 November 2000 14 April were considered nonbreeding-season records. Observer(s) R. E. Martin (Sheyenne Lake Breeding Bird Survey) (Berkey 1993) IGL AND MARTIN: MOCKINGBIRD OCCURRENCES IN NORTH DAKOTA Season* Breeding D. Griffiths (Bentley Breeding Breeding Bird Survey) (Berkey 1993) E. M. Madden (Berkey 1993) Bismarck-Mandan Christmas Bird Count (LeBaron 1994) D. Griffiths and C. Griffiths (Martin 1994) G. B. Berkey (Martin 1994) R. E. Martin (Martin 1994) D. O. Lambeth (Martin 1994) H. C. Talkington R. E. Martin D. Griffiths and C. Griffiths L. D. Igl, D. H. Johnson, and J. T. Price K. Askerooth R. Ziarno J. Hoff (Berkey 1995) R. N. Randall B. Boyd C. RR tama D. Wiesenborn H. A. Kantrud C. R. Gjervold and R. Gjervold 7. Hat +s H. C. Talkington J. Maxwell and B. Maxwell D. Griffiths (Bentley Breeding Bird Survey) L. D. Ig] (Martin 1998) J. Lefor C. D. Ellingson D. Griffiths and C. Griffiths M. A. Sovada and R. J. Greenwood: also reported on Jamestown Christmas Bird Count during Count week (Randall 2000) C. Norheim Breeding Nonbreeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Nonbreeding Breeding Breeding Breeding Nonbreeding Nonbreeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Breeding Nonbreeding Nonbreeding Breeding Nonbreeding Nonbreeding Number of birds one two + three fledglings one one one one one one one one one one two + nest (with four eggs) one one one one one 97 Diets of Northern Flying Squirrels, Glaucomys sabrinus, in Southeast Alaska SANJAY PYARE!, WINSTON P. SMITH2, JEFFREY V. NICHOLLS”, and JosEPH A. CooK?*+ ‘Denver Zoological Foundation, P.O. Box 20377, Juneau, Alaska 99802 USA [corresponding author] 2United States Forest Service, Pacific Northwest Region, Forestry Sciences Laboratory, 2770 Sherwood Lane, Suite 2A, Juneau, Alaska 99801 USA 3University of Alaska Museum, Fairbanks, Alaska 99775-6960 USA 4Current address: Department of Biological Sciences, Idaho State University, Pocatello, Idaho 83209-8007 USA Sanjay Pyare, Winston P. Smith, Jeffrey V. Nicholls, and Joseph A. Cook. 2002. Diets of Northern Flying Squirrels, Glaucomys sabrinus, in southeast Alaska. Canadian Field-Naturalist 116(1): 98-103. We examined the diet of the Northern Flying Squirrel (Glaucomys sabrinus) during summer and autumn seasons in tem- perate rain-forest habitat of Southeast Alaska, a region in which the ecology of this species is poorly understood. Truffles, a food item that is commonly consumed by squirrels during snow-free periods outside of Alaska, were present less frequent- ly in squirrel feces than two other food items, epigeous fungi and vegetation, although no food item dominated fecal com- position. Truffles were less frequent in fecal samples from mixed-conifer muskeg habitats than from old-growth forest habitats. Overall, we found that squirrels consumed a total of five truffle genera; Elaphomyces and Hymenogaster being the most common. Compared to populations in the western contiguous United States, squirrel populations in Southeast Alaska consumed truffles less frequently and consumed a smaller total number of truffle genera. In addition, samples from individ- ual squirrels in Alaska tended to contain fewer genera than samples from the contiguous United States. Finally, squirrels in Alaska consumed other food items such as vascular vegetation, lichens, and mushrooms more frequently than squirrels in other geographic areas. These patterns suggest that the association between flying squirrels and truffles may be relatively weaker in Southeast Alaska than has been documented elsewhere. Consequently, additional information on life history and ecology of flying squirrels is warranted before forest management guidelines can be developed. Key Words: Alaska, diet, lichens, mycophagy, Northern Flying Squirrels, old-growth, Pacific Northwest, truffles Northern Flying Squirrels (Glaucomys sabrinus) abundant and where fungal composition is relatively are mycophagous and frequently feed on the fruiting well documented, particularly in southern regions of bodies of hypogeous fungi (i.e., truffles) in conifer- the Pacific Northwest, such as in northern California, ous forests of western North America (Carey et al. Oregon, and Washington, U.S.A. (Carey et al. 2000; 2000; Colgan et al. 1997; Hall 1991; Maser et al. Colgan et al. 1997; Hall 1991; Maser et al. 1985; 1985; Maser et al. 1986; Maser and Maser 1988; Maser et al. 1986; Maser and Maser 1988; Pyare and Pyare and Longland 2001; Rosentreter et al. 1997; Longland 2001; Waters and Zabel 1995). In contrast, Waters and Zabel 1995). Because spores of these in northern regions of the Pacific Northwest, such as fungi are mycorrhizal and remain viable after pas- Southeast Alaska and adjacent British Columbia, lit- sage through small mammal digestive tracts, flying _ tle is known about the ecology and food habits of fly- squirrels are thought to play an important ecological ing squirrels, nor is much known about the distribu- role by dispersing these spores (Cazares and Trappe _ tion of truffles. This is true despite that coastal and 1990; Cork and Kenagy 1989; Kotter and Farentinos interior forests of this region provide extensive habi- 1984; Li et al. 1986; Maser and Maser 1988). tat for flying squirrels, and provide an abundance of During winter, truffles are uncommon food items conifers that potentially serve as ectomycorrhizal for flying squirrels because truffles are less abundant hosts for truffle fungi. In general, however, it is and/or snow cover may limit access to these below- thought that truffles are relatively uncommon in ground fungi. Thus, alternative and readily available northern forests possibly because climatic conditions food items, such as lichens, are predominant winter may be too extreme and/or this region may simply food items for flying squirrels. During snow-free _ represent the northern limits of truffle distribution for periods, however, such as in summer and autumn, some species (E. Cazares, personal communication). flying squirrels are thought to specialize on truffles | Thus, even during snow-free periods, truffles may be because flying-squirrel stomachs and feces generally — less common in this region, and consequently, flying contain a diverse array of truffle genera. Furthermore, squirrels may be more likely to exhibit generalist- squirrels exhibit preferences for truffles over other type feeding behavior there (Thysell et a. 1997). food items (Zabel and Waters 1997). For the most » Basic information about squirrel ecology and food part, however, our knowledge about these squirrels is habits in this region would have practical merit for restricted to geographic areas where these fungi are contemporary objectives regarding the conservation 98 2002 Alexander Archipelago ~ i “9 on a ond AR A Canada Prince of _ es Island Complex FiGurE 1. Location of study area (Prince of Wales Island complex) in the southern Alexander Archipelago of Southeast Alaska, U.S.A. Numbered locations are as follows: (1) Prince of Wales Island, (2) Heceta Island, (3) El Capitain Island, (4) Tuxekan Island, and (5) North Island. of the flying squirrel. In addition, an endemic sub- species, the Prince of Wales Northern Flying Squirrel (G. s. griseifrons), is found in Southeast Alaska (Demboski et al. 1998; McDonald and Cook 1996). Thus, to provide a more comprehensive understanding of flying-squirrel ecology, we evaluat- ed diet patterns of flying squirrels in temperate rain- forest habitats of Southeast Alaska during snow-free periods and compared these patterns to those in the western contiguous United States. Methods and Materials We conducted microhistological analyses of fly- ing-squirrel feces collected from a total of 13 forest stands in the Prince of Wales Island complex in the southern Alexander Archipelago of Alaska (Figure 1). These coastal study sites ranged from 200-800 m PYARE, SMITH, NICHOLLS, AND COOK: DIETS OF FLYING SQUIRRELS IN ALASKA 99 in elevation. Ten stands consisted of old-growth (un- logged) forest habitat, six of which were on Prince of Wales Island (stands separated by > 5 km): Dargun Point (55°54’44” N, 133°15'36” W), Honker 1 (55°20'48” N, 132°50’08” W), Honker 2 (55°21'02” N, 132°50'36), Honker 3 (55°19'54” N, 132°49'37” W), Naukati Bay (55°51’25” N, 133°10’02” W) and Tuxekan Village (55°54’06” N, 133°14’55” W). An additional four old-growth stands were on El Capitan Island (55°55’59” N, 133°18'33” W), Heceta Island (55°46'00"N, 133°27'00"W), North Island (55°57'00" N, 133°19'00” W), and Tuxekan Island (55°51'11” N, 133°13'39” W). Finally, three stands consisting of mixed-conifer muskeg habitat were on Prince of Wales Island (stands separated > 5 km): Ball’s Lake (55°43’00" N, 132°49’00” W), Control Lake 1 (55°43'48"N, 132°48'54” W), and Control Lake 2 (55°43’24", 13249'39” W). Old-growth habitat included Sitka Spruce (Picea sitchensis), Western Hemlock (Tsuga heterophylla) and Red (Thuja plicata) and Yellow cedar (Chamaecyparis nootkatensis). Large trees, downed and decaying wood, snags and small gaps created by individual blow-downs all were key components of this habitat. Dense patches of Blueberry (Vaccinium spp.) dominated the understory. In contrast, mixed- conifer muskeg habitat exhibited poor drainage, shal- low organic soils, and only patches of mixed conifer vegetation that occurred on gently sloping lowlands and floodplains. Conifer vegetation included Yellow Cedar, Red Cedar, Western Hemlock, Mountain Hemlock (T. mertensiana), Shore Pine (Pinus contor- ta var. contorta), and Sitka Spruce. Understory vege- tation varied considerably within this habitat type. Open areas with little overstory consisted of a mix- ture of herbaceous species (sedges, grasses, Skunk Cabbage [Lysichitum americanum]) and Labrador Tea (Ledum glandulosum), whereas Blueberry domi- nated areas with well-developed overstories. Because muskegs are a previously undocumented habitat for Northern Flying Squirrels, we treated data from these stands separately. We collected fecal samples from flying squirrels in these habitats only during snow-free periods; 1.e., when truffles were most likely to be available and/or snow could not preclude access to truffles. On three of the old-growth sites (Honker 1-3) and all three of the muskeg sites, fecal samples were collected from the anus of individually-marked squirrels that were live-trapped in Tomahawk live-traps during autumn (September-October) of 1999 and 2000. No more than one sample was collected from each individual. We did not collect samples that appeared to consist of bait (a peanut butter-rolled oats mixture). For the remaining seven old-growth sites, feces were collect- ed from the digestive tract of museum specimens (University of Alaska Museum, Fairbanks, Alaska). These samples were originally collected during sum- mer (June-July) and autumn 1999. 100 THE CANADIAN FIELD-NATURALIST, Vol. 116 TABLE 1. Prevalence of different food items in fecal samples collected from flying squirrels in old-growth stands (n = 10) and muskeg stands (3) in Southeast Alaska. Truffle genera found in each stand are shown at right. % Occurrence N_ Truffles Lichen Vegetation Mushrooms _ Insect Truffle Genera Present Old Growth Stand Honker | 34 50 50 50 59 Pa Elaphomyces, Hymenogaster Honker 2 14 50 20 23 30 0 Sclerogaster Honker 3 zl 100 20 56 20 0 Elaphomyces Dargan Point 6 50 12 50 0 0 Hymenogaster Naukati 5 64 20 75 48 - Elaphomyces, Hymenogaster Tuxekan Village 10 20 64 24 40 0 Elaphomyces Heceta Island 17 40 10 64 23 12 Elaphomyces, Hymenogaster Tuxekan Island 26 36 36 60 24 12 Elaphomyces North Island 3 12 30 84 Be 4 Elaphomyces El Capitan Island 14 82 8 56 84 0 Hymenogaster xX 50.4 27 55.2 36.1 4.4 s.d 26.8 18.4 20.1 25.3 4.6 Muskeg Stands Ball’s Lake 1 4 + 20 60 50 0 Elaphomyces Control Lake 1 a 12 40 28 40 0 Elaphomyces Control Lake 2 8 40 0 50 88 0 Elaphomyces x 18.7 20 46 593 0.0 s.d 15:5 16.3 12:5 21.6 0.0 All samples were stored separately in glass vials with 70% ethanol. For analysis, small portions of each sample were mixed with 3ml of distilled H,0, and mixed vigorously together to achieve a homoge- neous solution. A small drop of the solution was mixed with a drop of Melzer’s reagent and placed on a slide, covered with a 18 mm circular slide cover, and observed using bright-field microscopy at 40 x. We identified the following different food items in 25 randomly selected field-of-views: the spores of hypogeous fungi and epigeous fungi, algal cells from lichen, and remains of vascular vegeta- tion and insects. Spores of hypogeous fungi were further identified to genus using a spore key (Castellano et al. 1989). Results We analyzed a total of 151 fecal samples from old-growth forest sites, and 17 samples from muskeg sites. Among old-growth samples, vascular vegeta- tion was the most frequent food item (x + se = 55 + 6.4%), while truffle spores were second most frequent (50 + 8.5%) (Table 1). In contrast, among muskeg samples, both epigeous fungi and vegetation items were more frequent (59+ 12.5% and 46 + 7.2%, respectively) than truffles (19 + 8.9%). TABLE 2. Truffle-consumption patterns of various flying-squirrel populations in western North America. All data were collected in mature conifer-forest habitats. Seasons correspond to the following months: Spring (March-May), Summer (June-August), Autumn (September-November), and Winter (December-February). Location and Source Nevada County, California: Hall 1991 Lake Tahoe Basin, California: Pyare and Longland 2001 Lassen County, California: McKeever 1960 Plumas County, California: Waters and Zabel 1995 Douglas County, Oregon: Maser et al. 1986 Douglas and Coos County, Oregon: Carey et al. 1999 Lane and Linn County, Oregon: Maser et al. 1985 Blue Mountain Province, Oregon: Maser et al. 1985 Watson Falls, Oregon: Cazares et al. 1999 Payette National Forest, Idaho: Rosentreter et al. 1997 Thurston County, Washington: Colgan et al. 1997 Southeast Alaska (this study) % Occurrence By Season N Spring Summer Autumn Winter 107 92 92 81 81 93 - 100 100 - 24 83 100 100 56 88 - 100 - - 162 97 93 99 98 158 TS - - - 28 - - 100 - 63 - - 100 - 12 - 94 100 - 200 ra 78 - i 12 - - - 85 [31 - 50 50 - 2002 100 @ 80 oO oO Shien Alaska (S) re) ONon- ° 3p Alaska =~ t i420 0 a e se 4 & ao w ge ro) ser =e Food Type FIGURE 2. Comparison of food-item occurrence in pooled samples between southeast Alaska (n= 10 stands) and outside Alaska (9). Insect frequencies were not reported in most studies and therefore left out of this comparison. All samples were collected in mature conifer-forest habitat during snow-free peri- ods. Bars represent standard error. Non-Alaska data from: Cazares et al. 1999; Hall 1991; Maser et al. 1985; Maser et al. 1986; McKeever 1960; Pyare and Longland 2001; Rosentreter et al. 1997; and Waters and Zabel 1995. The frequency of lichen in samples was roughly simi- lar between muskeg and old-growth sites (20 + 9.4% and 27 + 5.8%, respectively). Five truffle genera were present in old-growth samples. Elaphomyces was the only truffle genus present in muskeg samples. Examination of spring and autumn fecal-analysis data from 11 other forest sites throughout the western United States revealed that overall, the frequency of truffles in fecal samples from southeast Alaska was comparatively low (Table 2; Figure 2). In contrast, the frequencies of lichens, vascular plants, and epi- geous fungi were higher in Southeast Alaska than in the contiguous western United States, although these differences were not significant (Figure 2). Although sample size had a strong influence on the number of truffle genera present in samples collected throughout the western contiguous United States (Figure 3), Southeast Alaska appears to be atypical and does not fit this general correlation. Taxonomic richness of Alaska diets was much lower than other- wise would have been predicted (roughly 22 genera) for the observed Alaska sample size (n = 151; old- growth habitat only). On average, individual squirrel samples from Alaska contained fewer genera than samples from California and Oregon (Figure 4). Discussion Although flying squirrels are mycophagous throughout their range, dietary patterns of popula- PYARE, SMITH, NICHOLLS, AND COOK: DIETS OF FLYING SQUIRRELS IN ALASKA 101 tions in Southeast Alaska appeared to differ from those in the western contiguous United States during snow-free periods. Compared to their southern coun- terparts, at the population level, squirrels in Southeast Alaska consumed truffles less frequently and consumed a smaller total number of truffle gen- era. Furthermore, individual squirrel samples from Alaska contained fewer genera than southern fecal samples. These patterns suggest that flying squirrels may be less dependent on truffles in Southeast Alaska, even during seasons when potential avail- ability is greatest. This is consistent with other stud- ies that have shown that the species may in some instances exhibit generalist-type feeding behavior. For example, Thysell et al. (1997) observed wild squirrels feeding on other types of highly digestible food items such as seeds and berries, and conse- quently concluded that previous studies involving fecal-analyses have typically underestimated the importance of such food items. More recently, Claridge et al. (1999) concluded that hypogeous fungi were of moderate nutritional value to captive flying squirrels, which failed to gain body mass when fed solely the common fungal sporocarp Rhizopogon vinicolor. Additional support for the idea that flying squirrels may in some part of their range be only marginally dependent on truffles comes from the southern Appalachian Mountains, where truffle abundance is relatively low and only a few common truffle genera are found in fecal sam- ples (Loeb et al. 2000; Weig] et al. 1992). 35 30 nd 2 @ 25 er o . > O 20 ; o + pasion eh ale Me aes eal ail = wn ® EOL yas a SE Alaska 200 100 0 50 150 No. Fecal Samples Analyzed FIGURE 3. Relationship between sample size and total num- ber of truffle genera detected in fecal samples of flying-squirrel populations in western North America (r = 0.90 not including Alaska data). Non- Alaska data from: Carey et al. 1999; Cazares et al. 1999; Colgan et al. 1997; Hall 1991; Maser et al. 1985; Maser et al. 1986; Pyare and Longland 2001; and Rosentreter et al. 1997. —-) i) a No. Truffle Genera Per Sample ow 2 1 0 yo & oe o 2 < > > > = = Location FIGURE 4. Comparison of the mean number of truffle genera per diet sample (i.e. per individual) among four fly- ing-squirrel populations. All data were collected during snow-free periods in mature conifer-forest habitat. Sample sizes and data sources are as fol- lows: Alaska (this study, n = 151); California (Pyare and Longland 2001, 93); NE Oregon (Maser et al. 1985, 63); and NW Oregon (Maser et al. 1985, 28). Although we did not assay truffle abundance at these study sites, and virtually nothing is known about truffle distribution in Southeast Alaska, the simplest explanation for the disparity in squirrel diets between Southeast Alaska and elsewhere may be due to low truffle availability. Alternatively, snow cover could not have been a factor for the rel- atively low consumption rates of truffles observed in our study. Nor is it likely that the disparity is explained by differences in food preferences, although we cannot exclude this as a possibility. We never encountered truffles at these study sites, nor did we ever witness mycophagous animal dig- gings, clues that both scientists and hobbyists com- monly use to locate these subterranean fruiting bod- ies (Waters and Zabel 1995). In contrast, S. P. fre- quently observed evidence of the association between these animal diggings and truffles in mixed-conifer forest of the Sierra Nevada Range, California. In samples from both old-growth and muskeg habitats in Alaska, the preponderance of Elaphomyces spp. further suggests that fungal diversity may be limited perhaps due to adverse cli- matic conditions of this cool, extremely wet region. Elaphomyces spp. have a relatively thick ( 0.5 cm) outer peridium and are thought of as being among the hardiest of hypogeous fruiting bodies. We are particularly skeptical that truffles fruit to any sig- THE CANADIAN FIELD-NATURALIST. Vol. 116 nificant degree in the more open areas of muskeg habitats, the soil of which is often poorly drained and flooded. The frequency of lichen, vegetation, and mush- rooms were all higher in Southeast Alaska samples than samples from elsewhere, although differences were not significant. These other food items may be relatively more important than our simple fecal pel- let analysis revealed; volume estimates of stomach contents would certainly have provided a better understanding of ingestion rates of different food items. Of particular relevance to flying squirrels in this region may be the abundance of arboreal lichens, which achieve high biomass and diversity. In particular, Bryoria spp. and Usnea spp. are known to be consumed by flying squirrels (Rosentreter et al. 1997; Zabel and Waters 1997) and appear to be pre- ferred over other lichens by captive animals (S. Pyare, unpublished data). We commonly observed both of these lichens in these old-growth forest and muskeg habitats during our three-year study of fly- ing squirrels in Alaska. Conservation and management measures for fly- ing squirrels have conventionally emphasized the underlying importance of hypogeous fungi (Loeb et al. 2000; Maser and Maser 1988; Zabel and Waters 1997). Management plans in Southeast Alaska and in other northern regions, such as coastal and interior British Columbia, however, may further need to rec- ognize the complete array of food items consumed by these northern populations. For instance, in the western contiguous United States, among studies of the impact of disturbances, such as fire, clear cutting, and selective harvesting, on flying squirrel popula- tions, there have typically been concomitant efforts to assay the impacts on truffle abundance or diversi- ty, while the impacts of disturbance on other food types have not been evaluated (Amaranthus et al. 1994; Carey et al. 2000; Cazares et al. 1999; Colgan 1997; Waters et al. 1994; Waters and Zabel 1995). Management strategies in northern regions, however, may require a more pluralistic understanding of dis- turbance impacts on lichens, vascular plants, and mushrooms, as well as belowground fungi. Furthermore, because patterns of foraging, move- ment, and habitat use differ as a consequence of these different feeding habits, management strategies may require additional efforts to understand the unique ecological niche flying squirrels occupy in this particular region. Acknowledgments Aren Eddingsaas, John Frisch, Jeff Hayes, Meade Krosby, Kari Murabito, and Kevin White diligently helped obtain diet samples of flying squirrels in Southeast Alaska. Collin Gillin of Tufts University kindly furnished laboratory space and a microscope for diet analyses. Efren Cazares and Jim Trappe had oe ja 2002 PYARE, SMITH, NICHOLLS, AND COOK: DIETS OF FLYING SQUIRRELS IN ALASKA 103 previously provided their expertise with fungal iden- tification at Oregon State University, Corvallis. This study was supported by the University of Alaska Museum, Fairbanks, Alaska, and the USDA Forest Service’s Forestry Sciences Lab in Juneau, Alaska. S. Pyare was supported by the Wildlife Conservation Society and the Denver Zoological Foundation dur- ing drafting of this manuscript. Literature Cited Amaranthus, M., J. M. Trappe, L. Bednar, and D. Arthur. 1997. Hypogeous fungal production in mature Douglas-fir forest fragments and surrounding plantations and its relation to coarse woody debris and animal mycophagy. Canadian Journal of Forest Research 24: 2157-2165. Carey, A. B., J. Kershner, B. Biswell, and L. A. Dominguez de Toledo. 2000. Ecological scale and for- | est development: squirrels, dietary fungi, and vascular plants in managed and unmanaged forests. Wildlife Monographs 142: 1-71. Castellano, M., J. M. Trappe, Z. Maser, and C. Maser. 1989. Keys to the spores of hypogeous fungi of northern temperate forests with special reference to animal mycophagy. Mad River Press, Eureka, California. 186 pages. Cazares, E., D. L. Luoma, M. P. Amaranthus, C. L. Chambers, and J. F. Lehmkuhl. 1999. Interaction of fungal sporocarp production with small mammal abun- dance and diet in Douglas-fir stands of the southern Cascade Range. Northwest Science 73: 64~76. Cazares, E., and J. M. Trappe. 1990. Spore dispersal of ectomycorrhizal fungi on a glacier forefront by mammal mycophagy. Oecologia 86: 507-510. Claridge, A., J. M. Trappe, S. J. Cork, and D. L. Claridge. 1999. Mycophagy by small mammals in the coniferous forests of North America: nutritional value of sporocarps of Rhizopogon vinicolor, a common hypogeous fungus. Journal of Comparative Physiology B 169: 172-178. Colgan, W. 1997. Diversity, productivity, and mycophagy of hypogeous mycorrhizal fungi in a variably thinned Douglas-fir forest. Ph.D. dissertation. Oregon State University, Corvallis, Oregon. Colgan, W., A. B. Carey, and J. M. Trappe. 1997. A reli- able method of analyzing dietaries of mycophagous small mammals. Northwestern Naturalist 78: 65-69. Cork, S. J., and G. J. Kenagy, 1989. Rates of gut passage and retention of hypogeous fungal spores in two forest dwelling rodents. Journal of Mammalogy 70: 512-519. Demboski, J. R., B. K. Jacobsen, and J. A. Cook. 1998. Endemism in the Alexander Archipelago: an assessment of genetic variation in northern flying squirrels (Rodentia: Glaucomys sabrinus). Canadian Journal of Zoology 76: 1771-1777. Hall, D. S. 1991. Diet of the northern flying squirrel at Sagehen Creek, California. Journal of Mammalogy 72: 615-617. Kotter, M. M., and R. C. Farentinos. 1984. Formulation of ponderosa pine ectomycorrhizae after inoculation with feces of tassel-eared squirrels. Mycologia 76: 758-760. Li, C. Y., C. Maser, Z. Maser, and B. Caldwell. 1986. Role of three rodents in forest nitrogen fixation in west- ern Oregon: another aspect of mammalian-fungus-tree mutualism. Great Basin Naturalist 46: 411-414. Loeb, S. C., F. H. Tainter, and E. Cazares. 2000. Habitat associations of hypogeous fungi in the southern Appala- chians: Implications for the endangered northern flying squirrel (Glaucomys sabrinus coloratus). American Midland Naturalist 144: 286-296. Maser, C., and Z. Maser. 1988. Interactions among squir- rels, mycorrhizal fungi, and coniferous forests in Oregon. Great Basin Naturalist 48: 358-369. Maser, C., Z. Maser, J. W. Witt, and G. Hunt. 1986. The northern flying squirrel: a mycophagist in Oregon. Canadian Journal of Zoology 64: 2086-2089. Maser, Z., C. Maser, and J. M. Trappe. 1985. Food habits of the northern flying squirrel in Oregon. Canadian Journal of Zoology 63: 1084-1088. McDonald, S. O., and J. A. Cook. 1996. The land mammal fauna of southeast Alaska. Canadian Field-Naturalist 110: 571-598. McKeever, S. 1960. Food of the northern flying squirrel in northeastern California. Journal of Mammalogy 41: 270-271. Pyare, S., and W. S. Longland. 2001. Patterns of ectomyc- orrhizal fungi consumption in remnant old-growth forests of the Sierra Nevada. Journal of Mammalogy 82: 681-689. Rosentreter, R., G. D. Hayward, and M. Wicklow- Howard. 1997. Northern flying squirrel seasonal food habits in the interior conifer forests of central Idaho, U.S.A. Northwest Science 71: 97-101. Thysell, D. R., L. J. Villa, and A. B. Carey. 1997. Observations of northern flying squirrel feeding behav- ior: use of non-truffle food items. Northwestern Naturalist 78: 87-92. Waters, J. W., K.S. McKelvey, C. J. Zabel, and W. W. Oliver. 1994. The effect of thinning and broadcast burn- ing On sporocarp production of hypogeous fungi. Canadian Journal of Forest Research 24: 1516-1522. Waters, J. W., and C. J. Zabel. 1995. Northern flying squirrel densities in fir forests of northeastern California. Journal of Wildlife Management 59: 858-866. Weigl, P. D., T. W. Knowles, and A. C. Boynton. 1992. The distribution and ecology of the northern flying squirrel in the Southern Appalachians. U.S. Fish and Wildlife Service. 140 pages. Zabel, C. J., and J. R. Waters. 1997. Food preferences of captive northern flying squirrels from the Lassen National Forest in northeastern California. Northwest Science 71: 103-107. Received 18 April 2001 Accepted 18 March 2002 104 THE CANADIAN FIELD-NATURALIST. Vol. 116 Nesting Activities of an Eastern Spiny Softshell Turtle, Apalone spinifera CLAUDE DAIGLE!, PATRICK GALOIS2, and YVES CHAGNON? ISociété de la Faune et des Parcs du Québec, direction de la recherche sur la faune, 675 René-Lévesque est, Québec, Québec GIR 5V7 Canada ; e-mail: claude.daigle @fapaq.gouv.qc.ca 2Société d’Histoire Naturelle de la Vallée du Saint-Laurent, 21 125 chemin Sainte-Marie, Sainte-Anne-de-Bellevue, Québec H9X 3Y7 Canada; e-mail: pagalois @aei.ca 3Société de la Faune et des Parcs du Québec, direction régionale de la Montérégie, 201 place Charles-Lemoyne, Longueuil, Québec J4K 2T5 Canada Daigle, Claude, Patrick Galois, and Yves Chagnon. 2002. Nesting activities of an Eastern Spiny Softshell Turtle, Apalone spinifera. Canadian Field-Naturalist 116(1): 104-107. Une tortue-molle a épines a été suivi étroitement durant la période de ponte de 1998. Cette tortue a démontré un patron quotidien d’activité régulier avant la ponte. Elle consacrait ses matinées a s’exposer au soleil et a se déplacer dans un secteur restreint, ses aprés-midi 4 se déplacer sur des distances de 2 4 3 km, pour finalement s’arréter en fin d’aprés-midi alors que durant la soirée, elle semblait étre 4 la recherche d’un endroit propice pour déposer ses ceufs. Cette tortue-molle a épines a démontré la capacité de se déplacer 4 contre courant sur une distance de plus de 7 km, en trois jours, pour aller pondre. An Eastern Spiny Softshell Turtle was closely followed during the 1998 nesting season. It showed a rather consistent daily pattern of activity. The morning was spent basking and moving around in a small area, large movements (2-3 km) were made during the afternoon and the turtle would stop for the evening, acting like it was searching for a suitable nesting site. This softshell turtle moved 7 km upstream in three days for nesting. Key Words : Eastern Spiny Softshell Turtle, Apalone spinifera, nesting, movement, activity, Québec, Canada. The information reported here was collected in the course of a larger study designed to help the con- servation of a unique Québec population of Eastern Spiny Softshell Turtle (Apalone spinifera) (Galois, 1999*). Attempts were made to follow three females to their nesting sites, but we were successful with only one. This adult female (535) was captured 18 June 1997 in Riviére aux Brochets (Figure 1), a trib- utary of Lake Champlain (73°13'’W, 45°04’N) and equipped with a radio transmitter (frequency 155.535; Holohil System Ltd, Ottawa, Canada) pro- vided with a mortality option that produces a double bip when the turtle remains still for more than four hours. The softshell spent the winter of 1998 in the Missisquoi River near Swanton, Vermont, U.S.A. (73°13'W, 44°58'N), moved north across Lake Champlain during the first half of May, and up in the Riviere aux Brochets, where it spent the following three weeks in the vicinity of a small island, more than 25 km away from its wintering area. We fol- lowed 535 for most of the daylight hours during the few days before nesting and the day after nesting in June 1998. Although nesting activities had been described before (Eigenman 1896; Newman 1906: Evermann and Clark 1920; Minton 1972; Ernst et al. 1994), few individuals were involved in those obser- vations, and daily activities shortly before and after ovoposition has not been reported previously. 9 June, mostly sunny, 26°C At our arrival (10:15), 535 was basking on the sunny side of the river, approximately 20 cm from water, with two other softshell females basking a few meters away. They started moving at around 11:30, going in and out of the water. By 13:30, 535 had left the area and moved upstream throughout the afternoon. At 18:00, after swimming about 2 km, it stopped and remained in the area until we left at 20:00. 10 June, sunny, 22°C When we arrived at 5:30, 535 was in the same area as the previous evening. The signal became double at 9:30, indicating that the turtle had not moved for the last 4 hours. At 9:30, we spotted 535 basking on a stone in the river, approximately 5 m from the shore. It entered the water at 10:30 but remained in the area. By noon, it had started moving upstream. The river was about 30 cm deep and 30 m wide, and steep banks allowed us to often have a good look in the water. On many occasions we could see 535 passing by. It swam vigorously, its neck fully extended, pointing its nose out of the water every 5 to 10 m and sometimes stopping in this posi- tion for a few seconds. The softshell kept this pace for the whole afternoon. At 17:00, it stopped next to a gravel bar 5 m long but only 10 to 20 cm above the 104 2002 DAIGLE, GALOIS, AND CHAGNON: NESTING OF SOFTSHELL 105 73°02’00 73°06’00 73°04’00 e) | JUNE {{th boa) 5 Venise-en-Québec os NOTRE-DAME- DE-STANBRIDG Pointe Campbell Le ---------.-1./ i ..Le Patt Ruisseau se, w au e-<. JUNE t0t SAINT-PIERRE-DE- VERONNE-A-PIKE-RIVER tewreno"t cod li Cours eau Edwin} a, Ruissaau Louis-Rocheleau (Lac Champlain) 73°06'00 73°04’00 73°02'00 FiGURE 1. Movements of a female Eastern Spiny Softshell Turtle during the nesting season. 106 water level, almost 3 km from its point of departure at noon. The softshell came out of the water at 17:30, moving around on the gravel bar and rubbing its chin on the substrate every now and then. It re-entered the water 10 minutes later, but climbed back on it at 19:50, departing about 20 seconds later when dis- turbed by a man passing on the road 10 m away. At 20:10, it climbed once more on the gravel bar, but was disturbed a second time by human activities. The softshell was still in the area when we left at 21:00. 11 June, mostly sunny, 22°C We returned to the area at 6:00. Although not visi- ble, 535 was still around. At 8:30, the signal became double and remained so until 10:15. For the next hour, the turtle seemed to be moving around, but staying in the area. At 11:15, 535 walked on the grav- el bar in spite of the presence of six ducks. Again, the softshell seemed to be rubbing its chin on the:ground. By 11:30, it was back in the water and swam inten- sively upstream throughout the afternoon. It climbed on the sunny shoreline of the river at 15:50, approxi- mately 2.5 km away from its daily departure point. It moved in and out of the water on a 30 m stretch of the river, next to a sharp curve. At 16:05, 535 walked on land for the sixth time, at the other end of the curve, about a hundred meters upstream from where it had first gone on land, but only 20-30 m in straight line through forest. The softshell started the excava- tion at 16:15 at the edge of the vegetation, into gravel and flat pebbles ranging in size from 1 to 10 cm long. The first part of the digging motion was slow, but it ended vigorously and pebbles were thrown up to 2 m away. The excavation stopped at 16:35. After a few minutes without moving (probably laying), the turtle started filling the hole at 16:45. The softshell ran back into the water at 16:49, 44 minutes after coming out of the water. The nest was excavated 2 m from the water and approximately 1 m above the water level, in full late afternoon sunshine (northwestern slope). Numerous other signs of nesting activity were present in the vicinity. The softshell immediately started swimming upstream. We observed 535 at 20:00, 1 km upstream from where it nested, about 100 m below a dam in a small village. We left the area for the night. 12 June, partially sunny, 24°C When we arrived at 6:40, the turtle had moved downstream about 0.5 km from where we last saw it the night before. The signal was double but became single 10 minutes later, indicating that the turtle just started to move after a stop of at least four hours. The radio indicated that the softshell was moving downstream. It swam by its nest at 7:00. A large Snapping Turtle (Chelydra serpentina) was digging a few meter from where 535 had excavated its nest the night before. We saw the softshell at 8:40, and THE CANADIAN FIELD-NATURALIST. Vol. 116 then again at 10:15. It was swimming vigorously, in the same way it had been during the few days it spent moving up the river. We stopped tracking the turtle around 11:00 to check the softshell’s nest to confirmed that it had really laid eggs. We found eggs exactly where the turtle dug the night before. In order to avoid any disturbance in the incubation pro- cess, eggs were covered, without counting them, as soon as we discovered the first egg. We tracked the turtle during the whole afternoon and by 17:30, 535 was back in the area it left 9 June. Travelling down- stream, it took the turtle one day to cover the same distance that had required three days moving upstream. Swimming time summed up to 11 hours travelling downstream and 17 hours moving upstream. Discussion Most events observed while tracking this softshell turtle are comparable to what other authors have described about A. spinifera nesting habits. We were impressed by the last part of the digging strokes, but Minton (1972) also reported that softshells were vig- orously throwing the sand several meters away. Newman (1906) reported that nesting process usual- ly lasts about one hour and 535 completed the entire process in 44 minutes. Breckenridge (1960) reports almost exactly the same nesting observations from Minnesota as ours, in terms of date, time, duration and behaviour. According to Newman (1906) nest- ing can also occur at midday. What was most notable about our observations was the daily pattern of activity preceding nesting. The day seemed divided into 3 parts. Morning would be spent basking and moving around in one area. Around noon, the turtle would leave the area and spend the whole afternoon swimming vigorously. By the end of the afternoon, the turtle would stop and then seem to be looking around for a suitable nesting spot. Our observations raised the following questions: Did the turtle choose this specific nesting site because it did not find any other suitable site while travelling or was it going for this exact spot because it knew it from past nesting experiences or because she had hatched there? We know that 535 was in this part of the river during the 1997 nesting season but it was not observed in nesting behaviour. What was the influence of the area’s other nests in 535 choice? Smell seemed to have played a significant role in her search. We believe that this is what brought her out of the water the day it nested. She first moved on land at the beginning of a sharp curve probably smelling the nests at the other end of this curve, only a few meter away through forest. As reported by Fletcher (1996*), our observations indicated that Apalone spinifera can move considerable distance in a short period of time, exploring different potential nesting sites. This would suggest that confirmation 2002 of the nesting site of any specific turtle can only be achieved through direct observation of the laying process. Acknowledgment We thank Roger Bider, Joél Bonin, Lyne Bouthillier, Jocelyne Brisebois, Clément Lanthier, Martin Léveillé, Jacques Jutras, David Rodrigue and Louis-Mare Soyez who provided help in the capture and tracking of the softshells during the whole pro- ject. David Rodrigue made useful comments on the first draft of this manuscript. Funding for the project was provided by the Ministére de |’Environnement et de la Faune du Québec, Fondation de la faune du Québec, Plan d’ Action Saint-Laurent, and Société d’ Histoire Naturelle de la Vallée du Saint-Laurent. Document Cited (marqued * in text) Fletcher, M. 1996. Management of softshell turtle habi- tat, year 1, 1996. Upper Thames river Conservation Authority, London, Ontario. 23 pages. Galois, P. 1999. Recherche de sites de nidification de la tortue-molle a épines (Apalone spinifera) a la riviére aux Brochets et inventaires de l’espéce sur la riviére Riche- lieu et la riviére des Outaouais. Gouvernement du Qué- DAIGLE, GALOIS, AND CHAGNON: NESTING OF SOFTSHELL 107 bec, Faune et Parcs, Service de |’aménagement et de l’exploitation de la faune, Longueuil, Plan d’ intervention sur la tortue-molle a épines au Québec, rapport d’étape 1998, xii + 95 pages. Literature Cited Breckenridge, W. J. 1960. A Spiny Soft-shelled Turtle nest study. Herpetologica 16: 284-285. Eigenman, C. H. 1896. Testudinae of Turkey Lake, Indiana. Proceedings of the Indiana academy of science. 1895, 5: 262-264. Ernst, C. H., J. E. Lovich, and R. W. Barbour. 1994. Turtles of the United States and Canada. Smithsonian Institution Press, Washington and London. 578 pages. Evermann, B. W., and H. W. Clark. 1920. Lake Maxi- nkuckee, a physical and biological survey. The Depart- ment of Conservation, State of Indiana. Volume 1, 660 pages. Minton, S. A. 1972. Amphibians and reptiles of Indiana. Indiana Academy of Sciences Monograph 3 : 1-346. Newman, H. H. 1906. The habits of certain tortoises. The Journal of Comparative Neurology and Psychology 16 : 126-152. Received 25 April 2000 Accepted 28 January 2002 Long-distance Movements by Female White-footed Mice, Peromyscus leucopus, in Extensive Mixed-wood Forest THOMAS J. MAIER Northeastern Research Station, USDA Forest Service, Holdsworth Natural Resources Center, University of Massachusetts, Amherst, Massachusetts 01003-9285 USA Maier, Thomas J. 2002. Long-distance movements by female White-footed Mice, Peromyscus leucopus, in extensive mixed-wood forest. Canadian Field-Naturalist 116(1): 108-111. Two adult female White-footed Mice (Peromyscus leucopus) were recovered 14 730m and 6840 m from where they were originally captured and tagged in central Massachusetts. Similar combinations of factors throughout an extensive forested landscape, including poor acorn (Quercus spp.) crops, high population densities of mice, and the exclusive social behaviour of other female mice, may have been responsible for these long distance movements. Key Words: White-footed Mouse, Peromyscus leucopus, dispersal, distance, mixed-wood forest, movement, Quercus, small mammals, social dynamics, Massachusetts. Movement in space-time remains a little under- stood process for most organisms (Turchin 1998). Even less understood are long-distance movements (Koenig et al. 1996), such as those made by small mammals dispersing across landscapes (Zollner and Lima 1999; Andrzejewski et al. 2000; Bowman et al. 2001). Nonetheless, even sporadic long-distance movements by small mammals may profoundly affect species’ evolution, population dynamics, and community structure (Cockburn 1992; Allen et al. 1993; Berlow 1999). As such, recognition of the spa- tial scale of such movements and their proximal causes becomes integral to understanding certain wildlife requirements, the effects of habitat perturba- tion, and possible epidemiological risks (Dickman et al. 1995; Kozakiewicz and Szacki 1995; Calisher et al. 1999). Here I report long-distance movement events by two female White-footed Mice (Pero- myscus leucopus) in extensive, New England mixed- wood forest, and discuss these movements in relation to the associated environmental conditions. Observations One month after capture on a small mammal trap- ping grid within the Quabbin watershed in central Massachusetts (42°27'13.7"N, 72°21'03.7"W), a tagged White-footed Mouse was recaptured 14 730 m north of the original capture site. The adult female (21.2 g), apparently healthy and not exhibiting any signs of reproductive activity, was ear-tagged with a No. | Monel tag (National Band and Tag Co., Newport, Kentucky, USA) on initial capture, 15 October 1998, and subsequently recaptured twice more at the same site. Approximately 4 weeks later, this mouse was trapped within a residence. Its identi- ty was verified based upon gender and the fact that no other small mammal researchers working in the area were using tag numbers in this sequence; a necropsy was not performed. A second tagged White-footed Mouse was trapped at another resi- dence 750 m northeast of the same Quabbin trapping grid during the same time period. This mouse was disposed of before verification of its identity. In mid- April 1993, the desiccated remains of another tagged White-footed Mouse were found outside a third resi- dence 6840 m north of its original capture site. This adult female, when captured on another small mam- mal trapping grid within the Quabbin watershed (42°26'50.9"N, 72°19'29.8"W) on 12 August 1992 (its only capture), appeared healthy and did not exhibit any signs of reproductive activity. Its identity was verified as described above. Both trapping grids on which these mice were ini- tially captured were located within a fenced portion of the Quabbin watershed, with vehicle entry restricted by locked gates (this property is managed by the Metropolitan District Commission to provide water for the Boston municipal area). The landscape between the initial capture and final recovery sites consisted of contiguous, closed-canopy mixed-wood forest of a predominately Red Oak (Quercus rubra)- White Pine (Pinus strobus)-Red Maple (Acer rubrum) forest cover type (Eyre 1980) with scattered residen- tial and agricultural openings. For the two mice travel- ing the furthest, potential travel routes were bisected by a minimum of six light-duty forest roads, three small streams, a two-lane primary highway, and a powerline right-of-way. Geomorphological features run mostly north-south and range in elevation from 200-380 m. Distances, unless otherwise specified, were measured using a Rockwell PLGR+96 Global Positioning System receiver with a minimum horizon- tal accuracy of 10 m, and analyzed using ARC/INFO on a Sun SPARC station. Other environmental conditions characterizing the 1998 period included high White-footed Mouse den- 108 2002 sity at the Quabbin trapping site (136/ha) and adja- cent sites (58—89/ha) (T. Maier, unpublished data). Few mice were actively breeding at this time, but large numbers of juveniles were observed (T. Maier, unpublished data). In comparison, historic densities for these mice for similar local habitat at the same time of year have ranged from 1 to 126/ha, with a mean density of 27.8/ha, SD = 30.6 (9 years data; W. Healy, USDA Forest Service, personal communica- tion). Area acorn (Quercus spp.) crops yielded > 75% less than the previous year, with virtually the entire 1998 crop consumed by Acorn Weevil (Curculio spp.) larvae (R. Field, USGS Biological Resources, personal communication; W. Healy, personal com- munication). Local climatological statistics for a 44- day period in 1998 encompassing the potential travel of mice were a mean maximum temperature of 10.5° C; mean minimum temperature of -0.9° C; mean tem- perature of 4.9° C, and total precipitation of 24.3 mm, all as rain from five precipitation events (NWS*). Environmental conditions characterizing the 1992 period beginning in August similarly included high White-footed Mouse density at the Quabbin trapping site (81/ha) and adjacent sites (40—126/ha), and the absence of an acorn crop throughout the area, with mice populations dramatically declining by October of the same year (Elkinton et al. 1996). All popula- tion density estimates were generated using program CAPTURE (Otis et al. 1978; White et al. 1982). Discussion Although the distance covered by the furthest trav- eling White-footed Mouse reported here supersedes previous reports of long-distance movement by either homing or non-homing individuals of this species (2 km, Wegner and Henein 1991), the genus Peromyscus (3.2 km, Murie and Murie 1931), and the family Muridae (14 km, Dickman et al. 1995), travel over the 14.7 km tract during the period from initial to final capture was well within the potential capabili- ty of Peromyscus mice. Murie and Murie (1931) reported a Deer Mouse (Peromyscus maniculatus) returning over 3 km to its capture site in 2 days. Similarly, Calisher et al. (1999) reported an adult Deer Mouse homing 1200 m within 24 hrs. Dice and Hoslett (1950), comparing the performances of seven species of Peromyscus on activity wheels, reported a maximum 24-hr record the equivalent of 37 km of travel by an individual White-footed Mouse. Besides possessing the stamina to travel great distances in short amounts of time, these mice have additionally been observed to surmount various obstacles to travel (Sheppe 1965; Teferi and Millar 1993; Calisher et al. 1999) similar to those found in this observation. The possibility that these mice may have been transported via human conveyance cannot be totally discounted, but this seems unlikely given that the mice were ini- tially captured within a restricted area behind locked MAIER: LONG-DISTANCE MOVEMENTS BY MICE 109 gates. Additionally, personal interviews with the resi- dents who collected these mice revealed that their closest physical proximity consisted of one or two instances of a briefly parked vehicle (< 4h during daylight) approximately 2.5 km from where the 1998 mice were initially captured. Most dispersal (sensu Lidicker and Stenseth 1992) by small rodent species has been observed to occur over short distances (McShea and Madison 1992; but see Koenig et al. 1996), with longest distances associ- ated with travel by males for most species. What fac- tors may have contributed to long-distance travel by female White-footed Mice? First, the contiguous for- est was likely advantageous to long-distance move- ments, compared to a more fragmented landscape (Yahner 1983; Krohne and Hoch 1999). Second, food deprivation has resulted in increased activity by Pero- myscuS (Falls 1968 and references therein). Although Acom Weevil larvae infestations may represent a sig- nificant dietary supplement, they are unsuitable as a primary energy source compared to sound acorns (Semel and Andersen 1988; Steele et al. 1996) from autumn through spring (Kirkpatrick and Pekins 2002). Third, an increase in population density has been observed to increase the number of dispersing Peromyscus mice (Fairbairn 1978; Krohne et al. 1984). Fourth, resident P. leucopus females may espe- cially exclude conspecific female immigrants when resources decline (Metzgar 1971; Wolff 1989). Together, these factors suggest that the combination of high mouse density and low food supply through- out a large extensively-forested area may have created a social environment that inhibited local immigration, as posited by Stickel (1968) and Anderson (1989), thereby increasing the probability of long-distance travel by emigrating female mice. Other reports of long-distance movement by small mammals through large areas of continuous habitat have suggested high population densities as the incentive (Clark et al. 1988; Bowman et al. 1999). Nevertheless, as observed by Fairbairn (1978), such movement is not likely a sim- ple function of density or its fluctuation, but rather relates to individual behaviour within a social struc- ture along with changing resource patterns. Proximal factors leading to long-range movements are difficult to elucidate; this may be due to the inter- action of contributing factors, and perhaps, in part, because so few observations of such movements exist. Various studies have attempted to assess the spatial mobility of small mammals using arrays of traps (Krohne et al. 1984; Liro and Szacki 1994; Dickman et al. 1995; Andrzejewski et al. 2000; Bowman et al. 2001); yet, due to logistical and methodological con- straints, the probability of detection of marked ani- mals may be expected to decline exponentially with distance, resulting in the significant underestimation of long-distance movements (Koenig et al. 1996; Turchin 1998). The long-distance movements report- 110 THE CANADIAN FIELD-NATURALIST , ed here may eventually prove more common than pre- viously thought, requiring those studies modeling the spatial distribution of similar small mammals to adopt anew metric. Acknowledgments I particularly thank T. Cox, M. Delano, D. Small, and J. Starkey for reporting the collection of these mice. I also thank D. Clark for site information; D. Goodwin for analyzing distance data; R. Field and W. Healy for data on local small mammal density and acorn crops; G. Boettner, J. Bowe, Jr., R. Brooks, R. DeGraaf, K. Doyle, J. Hestbeck, D. King, J. Wolff, and P. Zollner for suggestions regarding the manuscript; S. Cooter for ethological instruction; and the Massachusetts Metropolitan District Commission, Division of Watershed Management, Quabbin Section, for access to their management areas. Documents Cited (marked with* in text) [NWS] National Weather Service, NOAA. Orange and Worcester, Massachusetts, USA. Available from: URL: http://tgsv.nws.noaa.gov/er/box/climate/ Literature Cited Allen, J. C., W. M. Schaffer, and D. Rosko. 1993. Chaos reduces species extinction by amplifying local popula- tion noise. Nature 364: 229-232. Anderson, P. K. 1989. Dispersal in rodents: a resident fit- ness hypothesis. Special Publication Number 9, Ameri- can Society of Mammalogists. Andrzejewski, R., J. Babinska-Werka, A. Liro, E. Owadowska, and J. Szacki. 2000. 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Society of American Foresters, Washing- ton, D:C. Fairbairn, D. J. 1978. Dispersal of deer mice, Pero- myscus maniculatus: proximal causes and effects on fit- ness. Oecologia 32: 171-193. Falls, J. B. 1968. Activity. Pages 543-570 in Biology of Peromyscus (Rodentia). Edited by J. A. King. Special Publication Number 2, American Society of Mam- malogists. Kirkpatrick, R. L., and P. J. Pekins. 2002. Nutritional value of acorns for wildlife. Pages 173-181 in Oak for- est ecosystems: ecology and management for wildlife. Edited by W.J. McShea and W. M. Healy. John Hopkins University Press, Baltimore, Maryland. Koenig, W. D., D. Van Vuren, and P. N. Hooge. 1996. Detectability, philopatry, and the distribution of disper- sal distances in vertebrates. Trends in Ecology and Evolution 11: 514-517. Kozakiewicz, M., and J. Szacki. 1995. Movements of small mammals in a landscape: patch restriction or nomadism? Pages 78-94 in Landscape approaches in mammalian ecology and conservation. Edited by W. Z. Lidicker, Jr. University of Minnesota Press, Minneapolis. Krohne, D. T., B. A. Dubbs, and R. Baccus. 1984. An analysis of dispersal in an unmanipulated population of Peromyscus leucopus. American Midland Naturalist 112: 146-156. Krohne, D. T., and G. A. Hoch. 1999. Demography of Peromyscus leucopus populations on habitat patches: the role of dispersal. Canadian Journal of Zoology 77: 1247-1253. Lidkicker, W. Z., Jr., and N. C. Stenseth. 1992. To dis- perse or not to disperse: who does it and why? Pages 21-36 in Animal dispersal: small mammals as a model. Edited by N.C. Stenseth and W. Z. Lidicker, Jr. Chap- man & Hall, London. Liro, A., and J. Szacki. 1994. Movements of small mam- mals along two ecological corridors in suburban War- saw. Polish Ecological Studies 20: 227-231. McShea, W. J., and D. M. Madison. 1992. Alternative approaches to the study of small mammal dispersal: insights from radiotelemetry. Pages 319-332 in Animal dispersal: small mammals as a model. Edited by N.C. Stenseth and W. Z. Lidicker, Jr. Chapman & Hall, London. Metzgar, L. H. 1971. Behavioral population regulation in the woodmouse, Peromyscus leucopus. American Mid- land Naturalist 86: 434-448. ‘Murie, O. J., and A. Murie. 1931. Travels of Peromyscus. Journal of Mammalogy 12: 200-209. Otis, D. L., K. P. Burnham, G. C. White, and D. R. Anderson. 1978. Statistical inference from capture data on closed animal populations. Wildlife Monographs 62: 1-135. 2002 Semel, B., and D. C. Andersen. 1988. Vulnerability of acorn weevils (Coleoptera: Curculionidae) and attrac- tiveness of weevils and infested Quercus alba acorns to Peromyscus leucopus and Blarina brevicauda. American Midland Naturalist 119: 385-393. Sheppe, W. 1965. Dispersal by swimming in Peromyscus leucopus. Journal of Mammalogy 46: 336-337. Steele, M. A., L. Z. Hadj-Chikh, and J. Hazeltine. 1996. Caching and feeding decisions by Sciurus carolinensis: responses to weevil-infested acorns. Journal of Mammalogy 77: 305-314. Stickel, L. F. 1968. Home range and travels. Pages 373-411 in Biology of Peromyscus (Rodentia). Edited by J. A. King. Special Publication Number 2, American Society of Mammalogists. Teferi, T., and J. S. Millar. 1993. Long distance homing by the Deer Mouse, Peromyscus maniculatus. Canadian Field-Naturalist 107: 109-111. Turchin, P. 1998. Quantitative analysis of movement: measuring and modeling population redistribution in animals and plants. Sinauer Associates, Sunderland, Massachusetts. MAIER: LONG-DISTANCE MOVEMENTS BY MICE 111 Wegner, J., and K. Henein. 1991. Strategies for survival: white-footed mice and eastern chipmunks in an agricul- tural landscape. Page 90 in Proceedings of the World Congress of Landscape Ecology, held in Ottawa, Canada. International Association for Landscape Ecology. White, G. C., D. R. Anderson, K. P. Burnham, and D. L. Otis. 1982. Capture-recapture and removal methods for sampling closed populations. Los Alamos National Laboratory Technical Report LA-8787-NERP. Wolff, J. O. 1989. Social Behavior. Pages 271-291 in Advances in the study of Peromyscus (Rodentia). Edited by G.L. Kirkland, Jr. and J. N. Layne. Texas Tech University Press, Lubbock. Yahner, R. H. 1983. Population dynamics of small mam- mals in farmstead shelter belts. Journal of Mammalogy 64: 380-386. Zoliner, P. A., and S. L. Lima. 1999. Search strategies for landscape-level interpatch movements. Ecology 80: 1019-1030. Received 4 July 2000 Accepted 12 March 2002 Recent Trends in Stem Numbers in Goldenseal, Hydrastis canadensis, Populations at the Northern Limit of its Range ADRIANNE SINCLAIR! and PAUL M. CATLING? 'Box 214 Metcalfe, Ontario KOA 2P0 Canada; and Biology Department, University of Ottawa, Ottawa, Ontario KIN 6N5 Canada 2Bastern Cereal and Oilseed Research Centre, Biological Resources Program, Agriculture and Agri-food Canada, Research Branch, William Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada Sinclair, Adrianne, and Paul M. Catling. 2002. Recent trends in stem numbers in Goldenseal, Hydrastis canadensis, popu- lations at the northern limit of its range. Canadian Field-Naturalist 116(1): 112-115. A 15% increase in stem number occurred in 14 natural Ontario Goldenseal populations surveyed between 1998 and 2000, based on maps of stems in 1m? quadrats. Increase occurred in 10 of the 14 populations and was significant. The increase mostly involved small stems, but neither seedlings nor very small plants were observed suggesting that stem increase was largely due to vegetative reproduction by rhizomes and/or root tips. These data correspond to expectation since populations have not expanded rapidly nor spread substantially over the past few decades, but rather appear to have expanded slightly and slowly. Corresponding to earlier speculation, evidence is accumulating to suggest that substantial spreading and large population increases may be dependent on specific kinds and levels of disturbance some of which are no longer prevalent or frequent. Key Words: Goldenseal, Hydrastis canadensis, rare plant, medicinal plant, population trends, demography, population growth rate, population structure, management, monitoring, southwestern Ontario The top-selling and increasingly popular (Small and Catling 1999; Foster 2000*) deciduous wood- land plant, Goldenseal (Hydrastis canadensis L.) was recently re-evaluated as threatened in Canada (COSEWIC 2000*) due to few scattered popu- lations, habitat loss, and potential threat of over- collection for the medicinal plant market (White 1991*; Sinclair and Catling 2000a). The 20 popula- tions in Canada occur in southwestern Ontario at the northern limit of its native range, which extends from southern Wisconsin to southern Vermont and south to Arkansas and northern Georgia (Sinclair and Catling 2000a). The perennial rhizomes of Goldenseal, which produce an annual stem each year and enable vegetative reproduction, are col- lected for the medicinal plant market. The market demand for Goldenseal has been largely met by harvest of wild populations in the United States (Foster 1991; Small and Catling 1999; USFWS 1997*), which are now considered endangered, crit- ically imperilled, imperilled, rare, or uncommon, depending on the State (USFWS 1997*). Apart from anecdotal observations, information on growth trends in Goldenseal populations is lacking for all parts of its range. Such information is essen- tial for the development of effective conservation strategies and monitoring programs, and the need for it has been emphasized in the most recent review of goldenseal ecology and population biolo- gy (Gagnon 1999*). Here we present trends in pop- ulation structure and growth rate, over two years, from 14 natural Goldenseal populations in south- western Ontario. Methods In May of 1998, a 1m? quadrat was placed at the edge of a single colony within each of 14 naturally occurring Goldenseal populations. The 14 popula- tions occur in Lake Erie Lowland Ecoregion 135 and the southwestern portion of adjacent Manitoulin- Lake Simcoe Ecoregion 134 of the Mixedwoods Plains Ecozone (Ecological Stratification Working Group 1995). Site numbers in Figures 2, 3, and 4 correspond to locations (Sinclair and Catling 2000a) on file with the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) [see Sinclair and Catling 2000a and references therein]. Each quadrat was permanently marked with four alu- minum pins and a numbered aluminum tag. All stems were mapped within each quadrat according to the following size classes: small (single leaf < 5 inches in width); large (single leaf > 5 inches in width); and flowering (two or more leaves). The flowering stems are produced by larger rhizomes which are presumed to be the oldest. The smallest stems are generally associated with small rhizomes or root tips. Presence or absence of seedlings was noted. Due to the clonal nature of the plant, all of the stems may have devel- oped from a single individual. The distinction between ramets and genets was beyond the scope of the present survey. In May of 2000, the 14 quadrats were revisited and remapped according to the same size classes. In July of both years, quadrats were revisited to record fruiting stems. The quadrat maps developed in 2000 were compared to those developed in 1998 to document any change in stem numbers and size classes. Total number of stems, as well as 112 Number of Stems All B F ly eb Stem Class FIGURE 1. Histogram showing overall trend in population structure, from 1998 to 2000, based on a m? quadrat sample from each of 14 goldenseal populations in Ontario. B = fruiting, F = flowering, L = large non- flowering, LL = small non-flowering. numbers of small, large, flowering, and fruiting stems, over all sites, were compared between the two years using simple t-tests. Results and Discussion There was no evidence of harvest, or other recent disturbances (tree removal, trampling, etc.), at any of the 14 populations. Overall (14 quadrat samples com- bined), between 1998 and 2000, there was a signifi- cant increase of 15% (p = 0.0036, 550 to 635 stems, Figure 1). Ten out of 14 patches show an increase in the number of stems (Figure 2). The greater number of stems in 2000 is due to a significant increase in the number of small stems (p = 0.003, Figures 1 and 3). Almost all patches show an increase in small stems, and the increases are much greater than the losses (Figure 3). No seedlings or very small plants (< 1 inch leaf width) were noticed, suggesting that stems developed from roots or rhizomes of existing plants and that effective recruitment by seed was negligible. These data suggest that patches are slowly expanding by vegetative reproduction. Fruiting stems have also increased with marginal significance (p = 0.045, Figures 1 and 4). Only two patches show a decrease in the fruiting component (Figure 4). The increase in fruiting stems was not associated with increased flow- er production: virtually no change occurred in flower- ing stems (Figure 1). The increased number of fruit- ing stems in 2000 is therefore a result of a greater proportion of flowering stems producing fruit, which may have been a consequence of the weather. Although reduced herbivory could also be an expla- nation, there was no evidence for this. No increase in SINCLAIR AND CATLING: STEM NUMBERS IN GOLDENSEAL 113 30 20 10 Increase Decrease “2070 1B 235 6 7 11121415 171820 Site Number FIGURE 2. Increases or decreases in stem number, from 1998 to 2000, based on a m* quadrat sample from each of 14 goldenseal populations in Ontario. seed production capability actually occurred. Large stems have decreased but the loss is not statistically significant (Figure 1). These data correspond to expectation since popula- tions have not expanded rapidly nor spread substan- tially over the past few decades, but rather appear to have expanded slightly and slowly (Sinclair and Catling 2000a). The 15% increase in overall stem production and the increased fruit production were 24 16 Increase Decrease pe 1A 1B 23 5 671112141517 18 20 Site Number FIGURE 3. Increases or decreases in number of small, non- flowering stems, from 1998 to 2000, based on a m? quadrat sample from each of 14 Goldenseal popula- tions in Ontario. 114 24 16 o 8 w” 149] © oO = e QO---2=-------@------ wo 14] © ® QO -8 1A 1B 23567 111214151718 20 Site Number FIGURE 4. Increases or decreases in number of fruiting stems, from 1998 to 2000, based on a m? quadrat sample from each of 14 Goldenseal populations in Ontario. possibly a result of the relatively warmer summers of 1998 and 1999, which are characteristic of the main Goldenseal range further south. The lack of evidence for reproduction by seed sug- gests that a limitation must be operating on sexual reproduction other than pollination and dispersal (see Sinclair et al. 2000). The stability of Ontario Goldenseal patches correspond to trends in other woodland perennials, such as American Ginseng (Panax quinquefolius L.) and Wild Leek (Allium tric- occum Ait.), which are slow-growing with population growth rates near equilibrium (population stability or maintenance, Charron and Gagnon 1991; Nault and Gagnon 1993). Conversely, in disturbed habitats, where more resources such as light and nutrients are made available, plants tend to have highly variable population growth rates which can reach well above equilibrium (e.g., Furbish’s Lousewort (Pedicularis furbishiae S. Wats.) and Downy Oatgrass (Danthonia sericea Nutt.), as noted by (Gagnon 1999). Since occurrences of Goldenseal have been associated with disturbance (Sinclair and Catling 2000b), it seems probable that substantial spread and increase in Goldenseal populations is disturbance-dependent. The results of recently initiated field experiments, along with further population monitoring, will enable a bet- ter understanding of disturbance effects and provide useful information for recovery and management. Acknowledgments Fieldwork was supported by the Endangered Species Recovery Fund (ESRF) of World Wildlife Fund Canada and the Canadian Wildlife Service of THE CANADIAN FIELD-NATURALIST. Vol. 116 Environment Canada, and also by Agriculture and Agri-food Canada and the University of Ottawa. Keiko Lui, Josh Flanagan, and Bruce Sinclair provid- ed field assistance. Conservation Authorities and pri- vate landowners kindly allowed us to work on their properties. Documents Cited (marked * in text) COSEWIC (Committee on the Status of Endangered Wildlife in Canada). 2000. Canadian species at risk. May 2000. 28 pages. http:// www.cosewic.gc.ca/ COSEWIC/2000_list.pdf Foster, S. 2000. Goldenseal, Hydrastis canadensis: goldenseal’s future. Steven Foster Group. http://www. stevenfoster.com/education/monograph/golden- seal.html. Gagnon, D. 1999. A review of the ecology and population biology of goldenseal, and protocols for monitoring its populations. Final report to the Office of Scientific Authority of the US Fish and Wildlife Service. Groupe de recherche en écologie forestiére, Université du Québec a Montréal. http://www.nps.gov/plants/medicinal/pubs/ goldenseal. USFWS (United States Fish and Wildlife Service). 1997. Amendment to Appendix II of Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), United States of America - Proposal to Include Hydrastis canadensis in Appendix II, in accor- dance with Article 2, paragraph 2A. 31 pages. White, D. J. 1991. Status report on the Golden Seal, Hydrastis canadensis, in Canada. Committee on the Status of Endangered Wildlife in Canada, Ottawa. 21 pages. Literature Cited Charron, D., and D. Gagnon. 1991. The demography of northern populations of Panax quinquefolium (American ginseng). Journal of Ecology 79: 431-445. Ecological Stratification Working Group. 1995. A nation- al ecological framework for Canada. Agriculture and Agri-food Canada, Research Branch, Centre for Land and Biological Resources Research and Environment Canada, ~ State of the Environment Directorate, Ecozone Analysis Branch, Ottawa/Hull. 18 pages + appendix + map. Foster, S. 1991. Goldenseal Hydrastis canadensis. Botanical Series Number 309. American Botanical Council, Austin. 8 pages. Nault, A., and D. Gagnon. 1993. Ramet demography of Allium tricoccum, a spring ephemeral, perennial forest herb. Journal of Ecology 81: 101-119. Sinclair, A., and P. M. Catling. 2000a. Status of gold- enseal, Hydrastis canadensis (Ranunculaceae), in Canada. Canadian Field-Naturalist 114: 111-120. Sinclair, A. and P. M. Catling. 2000b. Ontario goldenseal populations in relation to habitat size, paths, and wood- land edges. Canadian Field-Naturalist 114(4): 652-655. Sinclair, A., P. M. Catling, and L. Dumouchel. 2000. Notes on the pollination and dispersal of goldenseal, Hydrastis canadensis L., in southwestern Ontario. Canadian Field-Naturalist 114: 499-501. Small, E., and P. M. Catling. 1999. Canadian medicinal crops. National Research Council of Canada, Ottawa. 240 pages. Received 6 February 2001 Accepted 12 March 2002 2002 Addenda Corrections and additional notes for “Ontario Goldenseal, Hydrastis canadensis, populations in relation to habitat size, paths, and woodland edges”. Canadian Field-Naturalist 114(4): 652-655. A substantial interest in this earlier article has lead to the following notes. One reader questioned whether or not paths in woodlands represent a random sample of the habitat? In other words paths may follow habitat subunits within wocdlands, such as higher ground. Thus certain plant distributions in a forest may be cor- related with paths but without the paths themselves being a causal factor. The woodlands containing popu- lations of Goldenseal that we examined were either flat or uniformly and gradually hummocky. The paths, which were associated with the use of tractors or all- terrain vehicles, appeared to be a random sample of the habitat. The paths were not conspicuously associat- ed with openings in the canopy, although some of the edges certainly received more light than the interior. Consequently disturbance apparently benefitting Goldenseal did not appear to be light-dependent, although increased light may contribute to a beneficial effect. Neither of the following two corrections changes the conclusions of the paper: (1) In calculating the effect of natural area size, all populations within a natural area should have been pooled. This applies to three cases (a) 11, 12, 13 and SINCLAIR AND CATLING: STEM NUMBERS IN GOLDENSEAL 115 14, (b) 16 and 17, (c) 1A and 1B, all in table 1. Instead of “a marginally significant decrease in num- ber of Goldenseal stems with increasing natural area size’, there was no significant relationship between number of Goldenseal stems and natural area size (P = 0.982, R?= 0.0). It is of interest to note here that when 3 large sites with large leverage were removed from the data set, there was still no significant rela- tionship with natural area size. (2) The figures in the D-value column in Table 1 are incorrect and so is the procedure described at the end of results in the first two sentences of the paragraph beginning “In order to ...”. This should read “In order to ... disturbance values were assigned to each site based on a scale of 1—5 (1 = undisturbed, 5 = heavily disturbed). The assessment was based sub- jectively on a number of factors including gaps in the canopy, evidence of a cutting operation (bulldoz- ing, stumps, log dragging, etc.), trampling and vehi- cle tracks, and presence of invading alien species and/or native species of more open sites.” Since this is actually what was done there is no change to the results or conclusions. The column of D values in Table 1 from top to bottom should be 3, 5, 2, 5, 1, 1, Did 25, eh Al. Bd. A aad: ADRIANNE SINCLAIR AND PAUL CATLING Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-food Canada, Research Branch, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada; e-mail: catlingp@em.agr.ca A Comparison of Techniques for Assessing Amphibian Assemblages on Streams in the Western Boreal Forest CyntuIA A. PAszKowskt!, GARRY SCRIMGEOUR2, BEVERLY A. GinGRAS!, and SHARON KENDALL” 'Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada; e-mail correspondence: cindy.paszkowski @ualberta.ca 2Forest Resources Business Unit, Alberta Research Council, P.O. Bag 4000, Vegreville, Alberta T9C 1T4 Canada Paszkowski, Cynthia A., Garry Scrimgeour, Beverly A. Gingras, and Sharon Kendall. 2002. A comparison of techniques for assessing amphibian assemblages on streams in the western boreal forest. Canadian Field-Naturalist 116(1): 116-119. The western boreal forest of Canada is rapidly being altered by agriculture, forestry, and energy extraction. As part of the Alberta Forest Biodiversity Monitoring Program’s effort to develop monitoring schemes for biota associated with streams, we compared the performance of four sampling techniques for amphibians (constrained visual searches, call surveys, pitfall traps, and above-ground funnel traps) in June through August along two low gradient streams. Of the four anuran amphib- ian species that occur in the region, we encountered adults and young-of-the-year of two, Wood Frog (Rana sylvatica) and Western Toad (Bufo boreas), with the former species dominating our surveys. Given the time and equipment constraints imposed by the monitoring-program protocol, visual surveys proved to be the most effective technique (88.5% of amphib- ian records) for simply determining the presence of species. Pitfall traps performed better than funnel traps. Call surveys were the least effective technique principally because sampling took place after most breeding was completed. Key Words: Alberta, biomonitoring, pitfall trap, visual survey, sampling techniques, Western Toad, Bufo boreas, Wood Frog, Rana sylvatica. Within the last decade the boreal forest in Alberta, which covers the northern half of the province (Rowe 1972), has become increasingly disturbed and fragmented by agricultural expansion, logging, ener- gy extraction, and road building. The cumulative effects of municipal and industrial activity on boreal ecosystem integrity and biodiversity are receiving increasing attention. The Alberta Forest Biodiversity Monitoring Program (AFBMP) was established in 1996 to identify a suite of quantifiable biological attributes (1.e., variables ranging from the individual to the community-level) to serve as indicators of sus- tainable forest management and to develop sampling protocols for detecting changes in terrestrial and aquatic biodiversity within Alberta’s 500 000 km? forested land base (Schneider1997*). AFBMP is based on a network of 1250 sites located on a systematic grid. Individual sites will be moni- tored once every 5 years with 250 sites visited annual- ly. Each site will include terrestrial habitat to be sam- pled for nonvascular and vascular plants, arthropods, birds, and mammals. The terrestrial habitat will be paired with a nearby stream location. Forest streams are sensitive to catchment disturbances and respond quickly to changes in adjacent habitats (Power et al. 1988; Welsh and Ollivier 1998). Because sites are numerous and remote (crews will typically fly in), sampling of all stream biota (benthic algae, macroin- vertebrates, fish) and habitat features will be complet- ' ed by a two-person crew in 2 - 4 days with only 0.5 - 1.5 days allotted to any particular taxonomic group. Sampling will occur in June, July, and August. Because of the presumed sensitivity of many species to degradation of both terrestrial and aquatic habitats (Green 1997), amphibians would seem a natural group for inclusion in AFBMP surveys. Likewise, the ambitious scope of the program offers a unique opportunity to increase our meager knowl- edge of the distribution and ecology of the six anu- ran and two caudate amphibians that occur in Alberta’s forests, particularly elucidating their use of streams and adjacent riparian habitat (Russell and Bauer 2000). However, given the time and resource constraints of the proposed survey design and well documented natural fluctuations in amphibian num- bers (Semlitsch 2000), two questions emerge: “What sampling techniques perform best?” and “Will sur- vey results support the use of amphibians as environ- mental indicators by AFBMP or other large-scale monitoring programs?’ To address these questions, our study compared four techniques for sampling amphibians (con- strained visual searches, call surveys, pitfall traps, and above-ground funnel traps) along two low gradi- ent streams in the boreal forest of northeastern Alberta. Surveys adopted the previously described constraints on effort, seasonal timing, and equipment prescribed by AFBMP stream protocols. Materials and Methods In 1999 amphibian assemblages were sampled along two third-order forest streams, Bear Creek (55°44’55"N, 112°12'16”W) and Crow Creek (55°37'26"N, 112°11'36”W), north of Wandering 116 2002 River, Alberta. Sites were within the known ranges of four species of amphibians (all anurans) that are widespread in boreal Alberta (Russell and Bauer 2000): Western Toad (Bufo boreas), Canadian Toad (Bufo hemiophrys), Boreal Chorus Frog. (Pseudacris maculata), and Wood Frog (Rana sylvatica). Visual surveys Amphibians were surveyed three times at each stream: June (14 - 20), July (12 - 16) and August (23 - 27). During a survey period, the two-person team performed low intensity area-constrained visual searches (Crump and Scott 1994) on two dates between 08:00 and 12:00 MDST. Searches were conducted along 200 m long x 1 m wide flagged transects, one on each side of the stream at a distance of 1 - 5 m from the streambed. One person walked slowly searching for animals active on the surface. Animals were captured by hand, identified to species, measured (snout-urostyle length, SUL), weighed, aged based on size and date of capture (“adults” = 1+year old, any individual captured in June or individuals > 27 mm SUL in July-August; young-of-the-year = newly metamorphosed individ- uals captured in July-August, < 27 mm SUL; based on C. Paszkowski, unpublished data for Wood Frog and Western Toad), then released. Forty-two percent of the individuals observed during surveys escaped capture. These animals were identified to species and assigned an age if possible. The location of each amphibian encountered (captured or observed) was recorded based on markers placed at 50-m intervals along the transect. The total time required to survey a transect ranged from 18 to 60 min. To control for effort, we calculated the total number of animals encountered per hour of searching. Call surveys At both sites, we performed call surveys with and without recording devices. Surveys with a “live” listener followed the audio strip technique (Zimmerman 1994). These were performed twice per month (one survey per date), at the same time and on the same 200 m transect as visual surveys. We used microcassette recorders (Panasonic RN- 202, Matsushita Electric Industrial Co., Ltd.) for fixed-position call surveys at each stream, twice each month, between 08:00 and 12:00. A recorder was placed on each side of the stream inside a plastic jug located 1.5 m up a tree. The two recorders were sep- arated by an additional 200 m measured linearly along the stream bank. Each recording session lasted 60 min, resulting in 4 h of recording per site each month. Trapping In June a team of two workers installed four pitfall arrays and six funnel-trap arrays at the two sites. Arrays were equally divided between the two stream banks, 2 - 10 m from the water, and were separated PASZKOWSKI, SCRIMGEOUR, GINGRAS, AND KENDALL: COMPARISON 117 by 50 m. Pitfall arrays consisted of four traps linked by drift fence, 30 cm high x 8 m long polyethylene strips attached to wooden stakes and anchored in the soil. Two traps were centrally located, on either side of the drift fence, and one trap was located at the end of either arm. Traps were buried plastic buckets, 24 cm deep and 24 cm wide, fitted with 7 mm? wire mesh funnels with 8 cm openings. Damp sponges were placed in each trap and twigs were inserted ver- tically to allow small mammals to escape. Arrays were left intact between monthly sessions, but traps were closed with plastic lids. Funnel-trap arrays consisted of two collapsible fiber-mesh minnow traps (50 cm long x 25 cm high x 25 cm wide with two inward directed openings 5.5 cm in diameter; Nylon Net, Memphis, Tennesse) centrally located on either side of a drift fence (as described above). Litter was used to partially cover traps and to create “ramps” to funnel openings. Damp sponges were placed inside. Funnel traps were removed between monthly sessions, but drift fences were left intact. During the three monthly sampling sessions, pit- fall and funnel traps were open for 3 - 6 consecutive days and checked daily before noon. Information on captured amphibians was recorded as described for searches. For pitfall and funnel traps, we calculated the number of animals caught per trap-night (1 trap- night = | trap opened for 24 h). Results Visual surveys were the most effective sampling technique, in terms of the combined number of cap- tures and observations (115), followed by pitfall arrays (10), funnel arrays (3) and live call surveys (2). Automated call surveys failed to record any amphibians. The two males heard calling were Western Toads at Crow Creek in June. Two Western Toads, one adult (37 mm SUL, pitfall trap) and one young-of-the-year (19 mm SUL, visual survey), were also captured at Crow Creek. All other amphib- ians encountered were Wood Frogs. The size distri- bution of Wood Frogs for all sampling combined was similar at the two sites: Bear Creek: mean SUL (mm) = 31.0 + 9.4 (SD), range: 20 - 52.5, N = 35; Crow Creek: mean SUL = 39.8 + 8.6, range: 20 - 59.5; N = 42. Most frogs captured were adults but we did record young-of-the-year (N = 19; 20 - 27 mm SUL) at both sites in July and August. Survey results differed between sites and among months. Encounter rates during visual surveys and capture rates with funnel traps were greater at Crow Creek (Figure la, b). Pitfall capture rates were greater at Bear Creek (Figure 1b). Visual survey encounter rates and funnel-trap capture rates gradu- ally decreased as the summer progressed (Figure Ic, d), whereas pitfall capture rates increased in August, reflecting captures of young-of-the year (Figure Id). 118 No. of encounters / h of visual survey Oo (= a So 7 0.040 0.030 0.020 0.010 No. of captures / total trap-nights 0.000 Crow Bear THE CANADIAN FIELD-NATURALIST. No. of captures / total trap-nights Vol. 116 hs NO c @ visual survey O pitfall traps — funnel traps = oO co No. of encounters / h of visual survey b a 0.030 0.020 0.010 0.000 August FiGurE 1. Encounter and capture rates for the Wood Frog at Bear Creek and Crow Creek using visual surveys (a), and pit- fall and funnel traps (b) for the three monthly sampling sessions combined. Monthly encounter and capture rates for the Wood Frog using visual surveys (c), and pitfall and funnel traps (d) for the two sites combined. Total time devoted to visual surveys was 8.05 h at Bear Creek and 8.12 h at Crow Creek. Trapping effort totaled 216 pitfall and 156 funnel trap-nights at Bear Creek, and 188 pitfall and 140 funnel trap-nights at Crow Creek. Discussion We documented substantial differences in the per- formance of the four sampling techniques. Live and automated call surveys largely failed. The proposed AFBMP schedule for sampling streams simply does not coincide with the breeding season of anuran amphibians in boreal Alberta. Calling begins in late April, peaks in May and only Bufo species (which we did detect) continue breeding into June (Russell and Bauer 2000). Although relatively quick and easy, call surveys make little biological sense in this program. Pitfall and funnel trapping sessions resulted in few amphibian records, accounting for only 10% of the total number of individuals. This was a disappointing performance considering that the two workers required 18 h and 13.5 h to install the full suite of pitfall and funnel-trap arrays, respectively. Using our pitfall trapping effort and design (16 traps in 4 arrays), two overnight trapping sessions were required to yield a 70% probability of capturing one Wood Frog. Funnel traps never even reached this level of performance. Trapping offers a reliable tech- nique at sites where low travel costs and easy access permit sampling over an extended time period, par- ticularly in upland habitats where amphibian densi- ties are relatively low or occurrence sporadic (Szaro et al. 1988). Pitfall trapping did result in the capture of newly metamorphosed Wood Frogs, thus provid- ing evidence of local breeding, possibly in Beaver (Castor canadensis) impoundments which were common on both streams. Visual searches required relatively small invest- ments in time and materials and proved to be the most successful technique for sampling streams. Wood Frogs were seen on 22 of 24 surveys. During these 22 surveys, searching along a transect for only 50 m offered a 70% probability of encountering at least one Wood Frog. Like pitfall trapping, visual searches encountered young-of-the year frogs. Visual surveys were well suited to the study streams because riparian areas were characterized by a low occurrence of woody debris and relatively sparse understory, features not shared by all forest streams in Alberta. Even our searches yielded fewer amphib- ians as the summer progressed and streamside grass- 2002 es and forbs increased in height and density. Thus, although constrained visual searches might best meet AFBMP criteria, data generated by a single visit to a location will be strongly influenced by habitat struc- ture, season, weather, time of day, and surveyor competence (Crump and Scott 1994). Our surveys encountered only two of four anuran amphibians that occur in northeastern Alberta (Russell and Bauer 2000). As Roberts and Lewin (1979) reported from visual surveys conducted 20 years earlier, the Wood Frog overwhelmingly domi- nated our records. The absence of the common, but cryptic, Boreal Chorus Frog can be attributed to sampling biases linked to season (nonbreeding ver- sus breeding and calling) and habitat (streams versus ponds). The paucity of toads may reflect insufficient sampling effort to detect these less abundant and more fossorial species. If AFMBP maintains its proposed sampling proto- cols and adopts constrained visual searches to moni- tor forest amphibians, our study suggests that the program will reliably document the presence/ absence of the Wood Frog at sites and collect some demographic information on poorly known western and northern populations (e.g., Leclair et al. 2000). It seems less likely that the program will be able to assess the health of boreal forest populations of amphibian species like the Canadian Toad which are of conservation concern in Alberta (Hamilton et al. 1998). In closing, we support amphibian surveys as part of innovative, large-scale projects such as the AFBMP. However, programs designed to sample simultaneously a large number of taxa over a large number of sites will inevitably result in compromises in terms of the types of data that can be collected for any given group of organisms and the accuracy of information concerning the distribution and status of species of interest. Acknowledgments We thank F. R. Cook, R. Seigel, and three anony- mous reviewers for providing comments on earlier drafts of this manuscript, and S. Boss for help in manuscript preparation. Documents Cited Schneider, R. 1997. Ecological diversity monitoring framework. Draft Discussion Paper, Alberta Forest Biodiversity Working Group, Edmonton, Alberta. www.fmf.ab.ca/p2.html. PASZKOWSKI, SCRIMGEOUR, GINGRAS, AND KENDALL: COMPARISON iY Literature Cited Crump, M. L., and N. J. Scott. 1994. Visual Encounter Surveys. Pages 84-92 in Measuring And Monitoring Biological Diversity: Standard Methods For Amphibians. Edited by W.R. Heyer, M.A. Donnelly, R. W. McDiarmid, L. C. Hayek, and M. S. Foster. Smithsonian Institute Press, Washington, D.C. Green, D. M. Editor. 1997. Amphibians in decline: Canadian studies of a global problem. Herpetological Conservation, Volume 1. Hamilton, I. M., J. L. Skilnick, H. Troughton, A. P. Russell, and G. L. Powell. 1998. Status of the Canadian Toad (Bufo hemiophrys) in Alberta. Alberta Environmental Protection, Wildlife Management Division, and the Alberta Conservation Association, Wildlife Status Report Number 12. Edmonton, Alberta. Leclair, R. Jr., M. H. Leclair, J. Dubios, and J. Daoust. 2000. Age and size of Wood Frogs, Rana sylvatica, from Kuujjuarapik, Northern Quebec. Canadian Field- Naturalist 114: 381-387. Power, M. E., R. J. Stout, C. E. Cushing, P. P., Harper, F. R. Hauser, W. J. Matthews, P. B. Moyle, B. Statzner, and I. R Wais De Badgen. 1988. Biotic and abiotic communities. Journal of the North American Benthological Society 7: 1-25. Roberts, W. and V. Lewin. 1979. Habitat utilization and population densities of the amphibians of northeastern Alberta. Canadian Field-Naturalist 93: 144-154. Rowe, J. F. 1972. Forest Regions of Canada. Canadian Forest Service 1300, Department of the Environment, Ottawa, Ontario. Russell, A. P., and A. M. Bauer. 2000. The Amphibians of Alberta. Second Edition. University of Calgary Press, Calgary, Alberta. Semlitsch, R. D. 2000. Principles for management of aquatic breeding amphibians. Journal of Wildlife Management 64: 615-631. Szaro, R. C., K. E. Severson, and D. R. Patton. Editors. 1988. Management of amphibians, reptiles and small mammals in North America. U.S. Department of Agriculture, Forest Service, General Technical Report RM-166. Welsh, H. H. Jr., and L. M Ollivier. 1998. Stream amphibians as indicators of ecosystem stress: a case study from California’s redwoods. Ecological Applications 8: 1118-1132. Zimmerman, B. L. 1994. Audio strip transects. Pages 92- 97 in Measuring And Monitoring Biological Diversity: Standard Methods For Amphibians. Edited by W.R. Heyer, M. A. Donnelly, R. W. McDiarmid, L. C. Hayek, and M.S. Foster. Smithsonian Institute Press, Washington, D.C. Received 16 March 2001 Accepted 26 February 2002 Notes A Rare Leucistic Spiny Dogfish, Squalus acanthias, from the Bay of Fundy, Nova Scotia BRIAN W. CoaAp! and JOHN GILHEN? \Canadian Museum of Nature, P. O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada 2Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Canada Coad, Brian W., and John Gilhen. 2002. A rare leucistic Spiny Dogfish, Squalus acanthias, from the Bay of Fundy, Nova Scotia. Canadian Field-Naturalist 116(1): 120-121. A rare leucistic specimen of the Spiny Dogfish, Squalus acanthias, is reported for the first time in Nova Scotia, Canada and North America. It is a 915 mm total length adult female bearing pups. This specimen is an overall yellowish colour but has a large, darkly pigmented area on the head dorsally, a spot at the first and second dorsal fin origins, and blotches on the caudal peduncle and in the pectoral axils. It was captured while dragging for flounder in 70 fathoms about 2 km off Boars Head, Bay of Fundy, Nova Scotia, Canada at 44° 23'N, 66° 15’W, on 22 May 1989 by Stewart Taylor, Captain of the long- liner Nadia C of Centreville, Digby County. Key Words: Spiny Dogfish, Squalus acanthias, leucism, first record, Bay of Fundy, Nova Scotia, Canada. On 22 May 1989, Stewart Taylor, Captain of the longliner Nadia C of Centreville, Digby County, was dragging for flounder in 70 fathoms, at a place known to fishermen as “Bear Cove Ground” about 2 km off Boars Head, Bay of Fundy, Nova Scotia, Canada at 44° 23'’N, 66° 15’W, when he also cap- tured a leucistic Spiny Dogfish, Squalis acanthias. It is a 915 mm total length adult female bearing pups, and is catalogued in the Nova Scotia Museum of Natural History as NSM85308. Spiny Dogfish are normally grey to black in over- all colour with conspicuous white spots on the flanks and a white belly. The Bay of Fundy specimen is an overall yellowish colour but has a large, darkly pig- mented area on the head dorsally, a spot at the first and second dorsal fin origins, and blotches on the caudal peduncle and in the pectoral axils (Figure 1). Albinism or leucism is rare in sharks, rays and their relatives although relatively common in bony fishes (Dawson 1964; 1966; 1971; Dawson and Heal 1976). Ben Brahim (1998) lists only 21 cases for 18 species in all the elasmobranchs. Additional records are given in Gopalan (1971) and Talent (1973) for a count of 24 cases for 20 species, 8 being total albi- nos, 12 being “partial” albinos (presumably leucis- tic), and 4 not clearly defined. Leucistic specimens have a paleness of colouration (Lincoln et al. 1982) with the eyes usually dark. The first record of a leucistic Spiny Dogfish in Canadian waters is there- fore of some rarity. Selected measurements on the specimen preserved in ethanol are as follows, with the methodology after Compagno (1984). Figures in parentheses are ranges for comparative material (NMC 66-600, 556 mm total length (TL), Nova Scotia, George’s Bank, ca. 42°00'N, 67°00'W; NMC 84-1512, 623 mm TL, off Sable Island, ca. 44°18’N, 59°45’W; NMC 68-19, 665-704 mm TL, Newfoundland, St. John’s, White’s Ledge, ca. 47°42'N, 52°42'W; NMC 67-9, 80 mm TL, New Brunswick, Saint John River, 45°22'N, 66°12'W). Prenarial length in preoral length 2.0 (1.8—2.0); narrowest mouth width in preoral length 1.5 (1.3-1.7); preorbital length in prebranchial length 2.6 (2.4—2.6); eye length in preorbital length 2.1 (1.9-2.3); first dorsal fin spine length in first dorsal fin height 1.4 (1.1—1.5); first dorsal fin height in fin length 2.1 (1.9-2.1); the second dorsal fin margin in first dorsal fin margin 1.3 (1.2—1.3). These morpho- metric characters fall within the ranges for normally coloured Spiny Dogfish and the leucistic specimen agrees with the description in Compagno (1984) in regards to other characters. The specimen is leucistic with the overall skin colouration being a light yellowish. All fins in the preserved specimen have a light brownish appear- ance, apparently internal and possibly an artefact of preservation. The dorsal fin spines are blackish at their bases. The eyes are black. There is a rectangu- lar patch on the dorsal head surface originating just posterior to the level of the spiracles and extending back over the pectoral fin. It becomes reticulated at the rear, and measures 71.6 mm in length and 24.5 mm in maximum width. An oval spot 5.9 mm long is at the first dorsal fin origin and a similar spot 5.6 mm long at the second dorsal fin origin. There is 120 2001 NOTES A FiGuRE 1. Dorso-lateral view of a rare leucistic Spiny Dogfish, Squalus acanthias, captured about 2 km off Boars Head, Bay of Fundy, Nova Scotia, Canada at 44°23'N, 66°15’W, by Stewart Taylor, Captain of the longliner Nadia C, of Centreville, Digby County, on 22 May 1989. an elongate blotch 9.7 mm long on the left caudal peduncle upper margin somewhat behind the caudal fin origin, matched by a similar blotch 10.0 mm long on the right side but somewhat posterior to the one on the left; and a blotch, about 15 mm wide, con- cealed in each pectoral fin axil. Frgiland (1975) provides the only other confirmed record of a leucistic Spiny Dogfish, taken in the North Sea in the late 1960s (with a verbal record of an earlier capture). This specimen was a female, 990 mm total length. Captain Taylor and other fishermen in the area of capture of the Nova Scotian specimen confirm that this is the first “pure white” dogfish seen. Normal-coloured dogfish were numerous in the area at the time of capture of the leucistic specimen. Acknowledgments We are indebted to Captain Taylor who captured this Spiny Dogfish, and to Claire Doucette, Fisheries Officer, Dibgy for bringing it to the Nova Scotia Museum. The photograph was taken by Ron Merrick, Media Services, Nova Scotia Museum. Literature Cited Ben Brahim, R. 1998. Albinisme chez une torpille ocel- lée, Torpedo (Torpedo) torpedo. Cybium 22: 83-86. Compagno, L. J. V. B. 1984. FAO Species Catalogue. Sharks of the World. An Annotated and illustrated cata- logue of shark species known to date. Food and Agriculture Organization, Rome, Fisheries Synopsis, 125, Volume 4: viii + 249 pages. Dawson, C. E. 1964. A bibliography of anomalies of fish- es. Gulf Research Reports 1: 308-399. Dawson, C. E. 1966. A bibliography of anomalies of fish- es. Supplement 1. Gulf Research Reports 2: 169-176. Dawson, C. E. 1971. A bibliography of anomalies of fish- es. Supplement 2. Gulf Research Reports 3: 215-239. Dawson, C. E., and E. Heal. 1976. A bibliography of anomalies of fishes. Supplement 3. Gulf Research Reports 5: 35-41. Frogiland, @. 1975. Albinisme hos hai [Albinism in sharks]. Fauna, Oslo 28: 170-173. Gopalan, U. K. 1971. On two abnormal sharks from Gujarat. Journal of the Bombay Natural History Society 68: 465-467. Lincoln, R. J., G. A. Boxshall, and P. F. Scott. 1982. A dictionary of ecology, evolution and systematics. Cambridge University Press, Cambridge. viii + 298 pages. Talent, L. G. 1973. Albinism in embryo gray smooth- hound sharks, Mustelus californicus, from Elkhorn Slough, Monterey Bay, California. Copeia 1973: 595- ae A Received 15 May 2000 Accepted 11 February 2002 Vol. 116 129 THE CANADIAN FIELD-N ATURALIST Anomalies in Incisor Wear of American Elk, Cervus elaphus, in the French River Delta, Ontario J. HAamre!, F. F. MALLory?, I. A. Fition!, G. S. BRown? and M. A. Jost? iNorthern Environmental Heritage Institute, Cambrian College of Applied Arts and Technology, 1400 Barrydowne Road, Sudbury, Ontario P3A 3V8 Canada 2Department of Biology, Laurentian University, Sudbury, Ontario P3E 2C6 Canada (Correspondence should be addressed to fmallory @nickel.laurentian.ca) Hamr, J., F. F. Mallory, I. A. Filion, G.S. Brown, and M. A. Jost. 2002. Anomalies in incisor wear of American Elk, Cervus elaphus, in the French River delta, Ontario. Canadian Field-Naturalist 116(1): 122-123. This article reports on limited observations of dental characteristics of American Elk introduced to the Burwash/French River area of Ontario from Alberta in the 1930s. Incisor wear was examined in 29 elk during a four-year study of two, geo- graphically distinct herds. All examined adult elk from the French River delta (six females and four males) had excessive front incisor (I-1, I-2) wear as compared to adult animals from the Burwash region (five females, one male). No wear was ascertained in calves and yearlings belonging to the French River herd. Radio-monitoring of winter foraging behaviour of elk in the French River delta revealed frequent use of rock tripe (Umbilicaria spp.), scraped from cliff faces along water- way shorelines. Key Words: American Elk, Cervus elaphus, incisor wear anomaly, Ontario. Historically, the eastern American Elk (Wapit1) inhabited most of southern Ontario and south-western Quebec. However, due to over-hunting and habitat degradation, elk were extirpated from the eastern half of the continent by the late 1800s (Bosveld 1996*). The small elk population residing between the north shore of Georgian Bay, Lake Huron and Sudbury, Ontario (46.00°N, 80.50°W) are remnant animals from a 1930s introduction of Rocky Mountain elk from Alberta (Polziehn et al. 1998). For at least 30 years, these Ontario elk have existed in two relatively distinct herds, one in the French River delta on Georgian Bay (50 km south of Sudbury), and the other in the vicinity of the former Burwash Industrial Farm, 20 km south of Sudbury (Ranta 1979; Brown 1998). This area is located within the Great Lakes-St. Lawrence ecotone and associated with mixed conifer and hardwood forests (Rowe 1972). The climate in this region is continental with some moderating influ- ences along Georgian Bay. As a part of a wider investigation of the remnant elk herds, 16 animals were captured and radio-col- lared between 1994 and 1997 to determine habitat use and causes of mortality. Field necropsies were performed on all deceased animals (collared and non-collared) that could be located during the study. In total, the incisors of 29 captured and dead animals were examined (Table 1). It was readily apparent that the incisor wear pat- tern differed between the two herds (Figures 1 and 2). The French River animals had severely worn lower incisors, especially I-1 and I-2. Animals between 6 and 15 years of age showed the highest amount of wear; however, no abrasion of lower incisors was apparent in calves nor yearlings. Elk replace their deciduous I-1 at 15 months of age and I-2 at 18 months (Bubenik 1982). Monitoring of winter foraging behaviour of instrumented animals revealed frequent scraping of lichens, particularly rock tripe, from steep cliff faces along the French River delta waterways. The use of lichens coincided with increased browsing on conifers during late win- ter (February-March). Among the consumed conifers were Common Juniper (Juniperus communis), Eastern White Cedar (Thuja occidentalis), and White Pine (Pinus strobus) (Jost et al. 1999). Animals from the Burwash herd also consumed conifers, but showed normal incisor wear. The variation in incisor wear appeared to result from differences in regional topography. The Burwash region had more rolling terrain, fewer cliff faces, and fewer waterways as compared to the French River delta. It is generally known that lichens comprise a large portion of the winter diet of Caribou and Reindeer (Rangifer tarandus) and are high in digestible carbo- hydrates and low in nitrogen (Thomas et al. 1984). Elk wintering in mature conifer forests in the Rocky Mountains readily consumed arboreal lichens (Hash 1973; Bohne 1974). Usnic acid is a common sec- ondary compound found in lichens (Culberson 1977) and data collected by Palo (1993) suggested that the addition of usnic acid to hay resulted in higher in TABLE 1. Number of examined animals in 3 age classes and 2 herds. calves yearlings adults Herd (O.5—1 year) (1.5—2 years) (2.5—15 years) Burwash 6 2 6 French River l 4 10 2002 Figure 1. Lower incisors of adult elk from the French River population, near Sudbury, Ontario showing severe wear associated with scraping lichens from cliff faces. vitro dry matter digestibility. Jenks and Leslie (1988) also found that in vitro digestion of White-tailed Deer (Odocoileus virginianus) forage was enhanced by lichens, increasing fermentation efficiency and forage digestibility. In the present study, use of lichens coincided with high snow depths and increased utilization of conifers during late winter (February and early March) and to a lesser extent early spring (late March and April). It is suggested that where available, rock tripe consumption enhanced the digestibility of conifers and other woody browse consumed by the studied elk in late winter and contributed to the observed variation in tooth wear patterns. Acknowledgments We thank the Rocky Mountain Elk Foundation, Safari Club International (Ontario Chapter), Tembec Forest Products, Ontario Federation of Anglers and Hunters, Ontario Ministry of Natural Resources, Department of National Defense, Parks Canada, Sudbury Fish and Game Protective Association, Cambrian College, and Laurentian University for pro- viding financial support. Heli North Aviation, Atwood Island Lodge, and Hartley Bay Marina, J. F. Zuchlinski, and T. L. Hillis, provided technical sup- port. Literature Cited Bosveld, H. J. 1996. A review of documented occur- rences of native elk (Cervus elaphus) in Ontario. Heritage Consultants, Parks Canada, Ontario Region, Cornwall, Ontario. 26 pages. NOTES 123 FIGURE 2. Lower incisors of adult elk from the Burwash population, showing a normal wear pattern. Brown, G.S. 1998. Spatial behaviour and habitat utiliza- tion by wapiti (Cervus elaphus) in the French River and Burwash regions of Ontario. M.Sc. thesis, Laurentian University, Sudbury. 77 pages. Bubenik, A. B. 1982. Physiology. Pages 125-179 in Elk of North America — Ecology and Management. Edited by J. W. Thomas and E. Toweill. Wildlife Management Institute, Harrisburg. 698 pages. Culberson, C. 1977. Chemical and botanical quick guide to lichen products. Bryologist 73: 77-377. Hash, H. S. 1973. Movements and food habits of the Lochsa elk. M.S. thesis, University of Idaho, Moscow. 85 pages. Jenks, J. A., and D. M. Leslie. 1988. Effect of lichen and in vitro methodology on digestibility of winter deer diets in Maine. Canadian Field-Naturalist 102: 216-220. Jost, M. A., J. Hamr, I. Filion, and F. F. Mallory. 1999. Forage selection by elk in habitats common to the French River/Burwash region of Ontario. Canadian Journal of Zoology 77: 1429-1438. Palo, R. T. 1993. Usnic acid, a secondary metabolite of lichens and its effect on in vitro digestibility in reindeer. Rangifer 13: 39-43. Polziehn, R. O., J. Hamr, F. F. Mallory, and C. Strobeck. 1998. Phylogenetic status of North American wapiti (Cervus elaphus) subspecies. Canadian Journal of Zoology 76: 998-1010. Ranta, B. 1979. Range and habitat relationships of wapiti (Cervus canadensis) in the Burwash-French River area of Ontario. M.Sc. thesis, Carleton University, Ottawa. 205 pages. Rowe, J. S. 1972. Forest regions of Canada. Publication number 1300. Canadian Forest Service, Ottawa. Thomas, D. C., P. Kroeger, and D. Herviux. 1984. /n vitro digestibilities of plants utilized by barren ground caribou. Arctic 37: 31-36. Received 24 March 2000 Accepted 4 April 2002 Vol. 116 124 THE CANADIAN FIELD-NATURALIST First record of the Hoary Bat, Lasiurus cinereus (Chiroptera: Vespertilionidae), from Prince Edward Island. DONALD F. McALPINE!, FRANCES MULDOON?, and ALEXANDER I. WANDELER2 \Natural Science Department, New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada; e-mail: dmcalpin@nb.aibn.com 2Canadian Food Inspection Agency, Animal Diseases Research Institute, 3851 Fallowfield Road, P.O. Box 11300, Station H, Nepean Ontario K2H 8P9 Canada. McAlpine, Donald F., Frances Muldoon, and Alexander I. Wandeler. 2002. First record of the Hoary Bat, Lasiurus cinereus (Chiroptera: Vespertilionidae), from Prince Edward Island. Canadian Field-Naturalist 116(1): 124—125. The Hoary Bat, Lasiurus cinereus, is reported from Prince Edward Island for the first time on the basis of an adult animal collected 17 August 1999 and submitted for rabies testing. Based on previous records for this species from the Atlantic region, the presence of the Hoary Bat on Prince Edward Island is not unexpected. Although Lasiurus cinereus appears to be rare in the province, bat surveys that might better establish the species status on the Island are lacking. Key Words: Hoary Bat, Lasiurus cinereus, new record, Prince Edward Island. The Hoary Bat, Lasiurus cinereus, is the most widespread of North American bats, and although infrequently encountered, is widely distributed in Canada (van Zyll de Jong 1985). Maunder (1988) summarized the 17 records for Atlantic Canada up to that time, including the first occurrence from New- foundland. Previously, Lasiurus cinereus has not been recorded on Prince Edward Island, and has been considered hypothetical for Kouchibouguac National Park, which borders the coast of New Brunswick adjacent to the Island (Maunder 1988; Tremblay 1992). However, recent acoustic surveys revealed the species presence in the park (H. Broders, personal communication to DFM). Based on these earlier reports from the region, the presence of Lasiurus cinereus on Prince Edward Island was not unexpected. The first specimen of the Hoary Bat on Prince Edward Island was collected 17 August 1999, at Charlottetown, Queens County, and submitted to the Canadian Food Inspection Agency for rabies testing. Although the rabies specimen submission form records that the animal was aggressive and was killed, it was negative for rabies virus. Only the head was submitted and it has now been deposited in the New Brunswick Museum collection (NBM 5801). The head includes the skull, with cranium slit, of an adult animal with pelage intact across the muzzle and the lower jaws posterior to the eyes. Nearly all records for the Hoary Bat in Atlantic Canada have been individuals encountered in late summer and autumn, with the Prince Edward Island record reported here following this pattern. Maunder (1988) suggested that late autumn tree bats may be present in the Atlantic region as a result of “drift migration” and van Zyll de Jong (1985) noted that this species has been recorded from widely scattered locations far beyond areas that appear to offer suit- able habitat. Although Lasiurus cinereus appears to be rare on Prince Edward Island, bat surveys that might better establish the species status on the Island are lacking. To date there is no evidence that the Hoary Bat breeds in the Atlantic region. Although there are two mid-July occurrences for Quebec (Pierre Aquin, per- sonal communication to DFM) there are only two July records for Maritime Canada, both for New Brunswick (12 and 29 July, NBM 1163, Tremblay 1989*). All other reports of the Hoary Bat from the Atlantic Provinces are from early August to 17 November. Acknowledgments For sharing current information on the status of the Hoary Bat in the Atlantic region with us we thank Pierre Aquin, Quebec Natural Heritage Data Centre; Kate Bredin, Atlantic Canada Conservation Data Centre; Hugh Broders, New Brunswick Cooperative Fish and Wildlife Research Unit, Uni- versity of New Brunswick; and Andrew Hebda, Nova Scotia Museum of Natural History. Documents Cited [marked* in text] Tremblay, E. 1989. A brief survey of the chiropterian fauna of Fundy National Park. Unpublished report, Parks Canada, Natural Resources Conservation, Fundy National Park, 19 pages plus appendices. Literature Cited Maunder, J. E. 1988. First Newfoundland record of the Hoary Bat, Lasiurus cinereus, with a discussion of other records of migratory tree bats in Atlantic Canada. Canadian Field-Naturalist 102: 726-728. Tremblay, E. 1992. Bats of Kouchibouguac and Fundy National Parks, New Brunswick, Canada. Pages 291-294 in Science and the Management of Protected Areas. Edited by J. Willison, S. Bondrup-Nielson, C. Drysdale, T. B. Herman, N. W. P. Munro, and T. L. Pollock. Elsevier, Amsterdam. 2002 van Zyll de Jong, C.G. 1985. Handbook of Canadian Mammals Number 2: Bats. National Museum of Natural Sciences, Ottawa, 212 pages. NOTES bs Received 30 May 2000 Accepted 24 January 2002 Wolf, Canis lupus, Response to Domestic Sled Dog, Canis familiaris, Activities in Central Yukon GERALD W. KuUZzyYKi-3 and KRISTIN M. KUZYK2 !Department of Renewable Resources, University of Alberta, Edmonton, Alberta T6G 2H1 Canada 2Faculty of Science, University of Alberta, Edmonton, Alberta T6G 2H1 Canada 3School of Environmental Sciences, Lakeland College, Vermilion, Alberta T9X 1K5 Canada Kuzyk, Gerald W., and Kristin M. Kuzyk. 2002. Wolf, Canis lupus, response to domestic sled dog, Canis familiaris, activi- ties in central Yukon. Canadian Field-Naturalist 116(1): 125-126. We present an observation of a lone Wolf (Canis lupus) regularly following teams of domestic sled dogs (Canis familiaris) and humans for distances up to 34 kilometers. The lone Wolf died of unknown causes, approximately 30 meters from a trapping trail used by sled dog teams and within vocalization distance of sled dogs at an occupied cabin. We also noted a pack of about seven Wolves that did not follow the same sled dog trail when they encountered it, and exchanged vocaliza- tions with the dogs. We suggest human-made trails can be especially important travel corridors for lone Wolves. Key Words: Wolf, Canis lupus, Dog, Canis familiaris, Black Bear, Ursus americanus, Lake Whitefish, Coregonus clupeaformis, travel corridor, Yukon, howling, communication. Wolves (Canis lupus) are ancestors of domestic dogs (Canis familiaris) (Morey 1992); are known to interbreed with dogs (Maagaard and Graugaard 1994); to aggressively interact with, and prey upon dogs (Tompa 1983); and to closely follow travelling sled dog teams (Karras 1975). Coppinger and Coppinger (1995) reviewed literature of livestock guard dogs in Europe and Asia, and applied this information to a study of Wolves and guard dogs in Minnesota, con- cluding that Wolves can treat dogs as conspecifics and not as prey. Reports of Wolf-dog interactions in the scientific literature are scarce (Fritts and Paul 1989), especially in areas of low human activity. In Yukon, most known Wolf-dog encounters are predatory, and usually occur in autumn and winter (Alan Baer, Yukon Fish and Wildlife Branch, personal communi- cation). From September 1991 to February 1992 we used 13 sled dogs for the purpose of trapping, south of Mayo in central Yukon (63° 30' N 135° 30' W). On 9 October 1991, at approximately 09:00 and 12:00 h, a pack of Wolves was heard howling within a couple kilometers of our trapline cabin. There was no response from the sled dogs that were chained sepa- rately within 40 meters of the trapline cabin. On 10 October a small Wolf track, presumably that of a pup, was noticed in the fresh snow within ten meters of a sled dog chained at the outside edge of the group of dogs. Two of the dogs had barked that night, but the cause may have been a Black Bear (Ursus americanus), as its tracks in fresh snow were noticed within 75 meters of the cabin. On 12 October, about two kilometers from the cabin, we discovered a Lake Whitefish (Coregonus clupea- formis) spawning run in a small narrows of less than five meters wide that was being used by a pack of Wolves and a Black Bear. Wolf and Black Bear tracks were obvious in the fresh snow near the creek. Whole and partially eaten Whitefish, as well as a cache of buried Whitefish, were found on the creek bank. The Wolves remained near the fish run for seven days until 19 October when the ice thickened and the fish run neared completion. On 8 December at 10:45 h, excited barking from the sled dogs drew our attention to a lone gray Wolf on the lake ice approximately 400 meters from our cabin. It quickly fled into the forest when it noticed humans near the cabin. The same afternoon, a single fresh Wolf track was seen on the main trapping trail, which had initially been made with snowshoes, and was regularly used for travel with sled dogs. A lone Wolf traveled on the trapping trail for approximately five kilometers and showed interest in a trap set for Wolverine (Gulo gulo) that had a Moose (Alces alces) head for bait. On the morning of 11 December a single large fresh Wolf track was noticed within 30 meters of a sled dog chained at the periphery of the group of dogs. A Wolf remained in the area of the cabin, successfully taking bait from traps and scav- enging cached Whitefish at the creek narrows for the next couple days. On 14 December a Wolf was caught in and then escaped from a #4 long spring 126 trap, set for Lynx (Lynx canadensis). The same evening from approximately 19:00—19:15 h, howling from a single Wolf was heard across the lake from the cabin, with no reply from the dogs. In late December and early January, lone Wolf tracks were a frequent occurrence on and near the main trapping trail. On the morning of 17 January 1992, we left the main cabin and traveled 17 kilome- ters to a smaller line cabin. During the night there was a light snowfall and the following morning, we noticed a single fresh Wolf track 30 meters from the cabin. A female sled dog in estrous was tied near the front door of the cabin, and no barking by the dogs was heard at night when the Wolf was nearby. On 19 January, during the return trip to the main cabin, we encountered a single fresh Wolf track in the snow, which suggested the same lone Wolf had followed us to the small line cabin. On 20 January, a fresh lone Wolf track was observed within two kilometers of the main cabin, suggesting the same individual Wolf had followed us on the return trip back to the main cabin, a distance totaling approximately 34 kilome- ters. Wolves maintain their territories by killing tres- passing Wolves (Mech 1991), so during this foray, the lone Wolf had traveled well inside an occupied Wolf pack territory and risked being killed by terri- torial Wolves. On 23 January at about 10:00 h, and approximate- ly three kilometers from the main cabin, excitement from the team of sled dogs alerted us to a dead Wolf in its bed about 30 meters from the trapping trail. We assume it was the same Wolf that had been in the locality, due to the gray pelage and the fact that fol- lowing this discovery, we saw no more evidence of lone Wolf tracks on the trails and heard no more howling from a single Wolf. The cause of the Wolf's death was undetermined; there were no signs of external injuries from other Wolves and rigor mortis had not occurred. The Wolf was an old male as determined by tooth wear, and was in an emaciated state, as indicated by the lack of body fat and pro- truding ribs, although the coat was still in prime con- dition. We believe the site of death for this Wolf was not random because it was only a short distance from the compacted trail the Wolf had used in recent trav- els. The site was also within vocalization distance of sled dogs and in view (three kilometers across a val- ley) of the occupied cabin. Human-made trails can provide easier movement for Wolves (James 1999) and as in this case, may be especially important for lone Wolves in poor condition. On 25 January 1992 a pack of an estimated seven Wolves encountered our trapping trail on a narrow creek, the same one used by the lone Wolf. There - was little snow at this location and the Wolves fanned out and did not follow the trail off the creek. On 2 February at approximately 12:00 h, while we were resting the two dog teams near the same loca- THE CANADIAN FIELD-NATURALIST Vol. 116 tion, a pack of Wolves began howling approximately five kilometers away. The air was still, the sky sunny and the temperature estimated at -20C. All the dogs became attentive, looked in the direction of the Wolves and began synchronously howling for approximately one minute. They then became silent and attentive, looking in the Wolves' direction. The Wolves and sled dogs exchanged howls once more, then there was prolonged silence and we left the area. Wolves howl as a means of territorial mainte- nance and as communication within and between packs (Harrington and Mech 1979). No scientific reference was found of Wolves and dogs communi- cating through howling, so this observation may help in further understanding Wolf-dog interactions. Acknowledgments We thank Bernard Menelon and Barry Pouche both of Mayo, Yukon, for allowing us the use of their trapline. Ludwig Carbyn (Canadian Wildlife Service, Edmonton, Alberta) and Alan Baer (Fish and Wildlife Branch, Whitehorse, Yukon) kindly reviewed this note and added helpful suggestions. Literature Cited Coppinger R., and L. Coppinger. 1995. Interactions between livestock guarding dogs and wolves. Pages 523-526. In Ecology and conservation of wolves in a changing world. Edited by L. N. Carbyn, S. H. Fritts and D.R. Seip. Canadian Circumpolar Institute, University of Alberta, Edmonton. Fritts, S. H., and W. J. Paul. 1989. Interactions of wolves and dogs in Minnesota. Wildlife Society Bulletin 17: 121-123. Harrington, F. H., and L. D. Mech. 1979. Wolf howling and its role in territory maintenance. Behaviour 68: 207-249. James, A. R. C. 1999. Effects of industrial development on the predator-prey relationship between wolves and caribou in northeastern Alberta. Ph.D. thesis, University of Alberta, Edmonton, Alberta. 70 pages. Karras, A. L. 1975. Face the north wind. Burns and Maceachern Limited, Don Mills, Ontario. 191 pages. Maagaard, L., and J. Graugaard. 1994. Female arctic wolf, Canis lupus arctos, mating with domestic dogs, Canis familiaris, in northeast Greenland. Canadian Field-Naturalist 108: 374~-375. Mech, L. D. 1991. The way of the wolf. Voyageur Press, Stillwater, Minnesota, 120 pages. Morey, D. F. 1992. Size, shape and development in the evolution of the domestic dog. Journal of Archeological Science 19: 181-204. Tompa. 1983. Problem wolf management in British Columbia: conflict and program evaluation. Pages 112- 119 in Wolves in Canada and Alaska: their status, biolo- gy, and management. Edited by L. N. Carbyn in Canadian Wildlife Service Report Series, Number 45. Received 10 June 2000 Accepted 22 February 2002 2002 NOTES i277 First Record of an Eastern Coyote, Canis latrans, in Labrador Tony E. Cuupss!, and FRANK R. PHILLIPS? ‘Department of National Defence, 5 Wing Goose Bay, Box 7002, Station A, Happy Valley-Goose Bay, Labrador, Newfoundland AOP 1S0, Canada "Department of Forest Resources and Agrifoods, Government of Newfoundland and Labrador, Box 3014, Station B, Happy Valley-Goose Bay, Labrador, Newfoundland AOP 1E0 Canada Chubbs, Tony E., and Frank R. Phillips. 2002. First record of an eastern Coyote, Canis latrans, in Labrador. Canadian Field-Naturalist 116(1): 127-129. An adult male Eastern Coyote, Canis latrans, trapped on 14 January 1995 along the Churchill River, is the first confirmed record for Labrador. This record is approximately 600 km north east of the previously accepted range limit in eastern Canada. Key Words: Eastern Coyote, Canis latrans, range, distribution, Labrador. Coyotes (Canis latrans) have been expanding their range northward and eastward in North America during the past century (Perkins and Mautz 1990; Thurber and Peterson 1991; Lehman et al. 1991; Parker 1995). Coyotes were first recorded in Québec in 1944 (Rand 1945) and had spread north across the Gaspé Peninsula by 1974 (Georges 1976). More recently, Coyotes have become established as a significant component of the boreal forests of southeastern Québec (Créte and Desrosiers 1995; Tremblay et al. 1998; Patterson et al. 1999). Coyotes immigrated to the island of Newfoundland on drift- ing pack ice and were first documented in 1985 (Parker 1995). Here, we report the first record of an Eastern Coyote in Labrador, approximately 600 km north east of the previously recorded range limit in Québec (George 1976; Parker 1995). On 14 January 1995, an adult male Coyote was trapped along the Churchill River (53°17'N, 60°15' W), approximately 2 km south east of Happy Valley—Goose Bay. The Coyote was estimated (F. Blake, personal communication) to weigh approxi- mately 15 kg. Although the weight could not be con- firmed, it is consistent with reported average weights of 13.5 kg, and 12.5 kg for adult male forest Coyotes in Québec (Tremblay et al. 1998; Poulle, M.-L. et al. 1995) and about half the weight (ca. 30 kg) of Wolves (Canis lupus) in Labrador (Parker and Luttich 1986). Dr. C. G. van Zyll de Jong of the National Museum of Natural Sciences, later exam- ined the skull and performed a quantitative compari- son of the Goose Bay skull with samples of Coyotes and Wolves of both sexes using a ratio diagram (Figure 1). Eleven different dimensions of the skull (see Nowak 1979) for each of the taxa and the Goose Bay specimen were compared. All measurements were converted into logarithms and the sample of male Wolves was used as the standard. The differ- ence (d) between the logarithmic value of the stan- dard and the other groups and the Goose Bay speci- men were calculated, plotted and connected by lines to aid in their interpretation (Figure 1). This compar- ison revealed that the Goose Bay skull ratio series was similar to those of the male and female Coyote samples and within two standard deviations from the male Coyote sample means. These results confirmed its identification as that of a large adult male Coyote (Dr. C. G. van Zyll de Jong, 17 December 1995, per- sonal communication). Recent information on expanding Coyote populations in eastern Québec also indicates that Coyotes appear to be larger in the eastern portion of their range (Thurber and Peterson 1991; Lariviére and Créte 1993). Although no previous records of Coyotes exist for Labrador, the possibility of an established population remains plausible, as they do coexist with Wolves in Alaska (Thurber and Peterson 1991), Alberta, British Columbia, and more recently Idaho (Pilgrim et al. 1998). No geographic barriers seem to exist to pre- vent Coyotes from extending their range northward into Labrador. Delineation of Coyote range expan- sion in Labrador may have been precluded by the fact that no systematic surveys have been conducted for any carnivore species in Labrador and low fur prices in the last decade have significantly decreased trapping effort resulting in only one specimen to date. The expansion of Coyotes into eastern forests has occurred due to high populations of White-tailed Deer (Odocoileus virginianus), high populations of humans and associated garbage, and changes in for- est communities, all of which have resulted in an improvement in food supply (Thurber and Peterson 1991; Tremblay et al. 1998). Both sedentary and migratory Caribou (Rangifer tarandus) herds tre- quent central and southern Labrador and Coyotes have been identified as a threat to remnant popula- tions of Woodland Caribou in southeastern Québec (Créte and Desrosiers 1995). Recent extensions of the winter range by migratory Caribou from the George River Caribou herd (Department of National Defence, unpublished data), and increased numbers of Moose (Alces alces) in southern Labrador 128 -0.3 —@®—C. Latrans (males, n=166) —#—C. latrans (females, n=111) —+— Goose Bay Specimen —O—-C. lupus (females, n=146) —t}—-C. lupus (males, n=233) - - @ - :Standard deviation C. lupus (male) - - + - :Standard deviation C. latrans (male) THE CANADIAN FIELD-NATURALIST Vol. 116 d = log x -log standard 10. LTB 11. HMO 13. DC 14. LP4 15. WM1 Skull Dimension FiGuRE 1. Ratio diagram comparing eleven standard cranial measurements of the Goose Bay, Labrador specimen with sam- ples of male and female Canis lupus and C. latrans. For a description of measurements and samples of C. Jupus and C. latrans see: Appendix B in Nowak (1979). (Chubbs and Schaefer 1997) may provide a suffi- cient prey and carrion base to support Coyotes through the winter. Coyotes are a highly adaptable species (Pilgrim et al. 1998) and are capable of remaining in an area with a seasonally reduced prey base and harsh weather conditions (Patterson et al. 1999). Although the boreal forest appears to be a rel- atively poor habitat for Coyotes (Tremblay et al. 1998), Coyotes can persist at low densities, depend- ing mainly on Snowshoe Hares (Lepus americanus), mice and voles (Patterson et al. 2000; M. Créte per- sonal communication). It has been known for some time (Lehman et al. 1991; Pilgrim et al. 1998) that hybridisation of Wolf and Coyote genotypes has already occurred in Québec due to the rapid north- east progression of Coyotes. Although this single record may not confirm a range extension, the potential for Coyote range expansion into Labrador appears to be plausible and even likely. The sparse human population of Labrador, lack of systematic surveys, and decreas- ing trapping effort may result in an existing sparse population going undetected. Despite the presence of a healthy Wolf population, which may limit or slow down colonisation through intraguild preda- tion, the increasingly human-transformed landscape may be the key to the establishment of Coyotes in Labrador. Acknowledgments We appreciate the support of the Newfoundland- Labrador Wildlife Division, Goose Bay, Labrador and J. A. Schaefer, who arranged for the skull mea- surements and identification conducted by the late C. G. van Zyll de Jong. Thanks are extended to trap- per Fred Blake for providing the skull and details of the capture location and to D. Laing for drafting the figure. Funding for this publication was provided through the Environmental Monitoring Program, through the Department of National Defence (DND). We thank DND (Major G. Humphries) for their con- tinued support of wildlife research in Labrador. We also thank M. Créte and two anonymous referees for commenting on an earlier draft of this manuscript. 2002 Literature Cited Chubbs, T. E., and J. A. Schaefer. 1997. Population growth of Moose, Alces alces, in Labrador. Canadian Field-Naturalist 111: 238-242. Créte, M., and A. Desrosiers. 1995. Range expansion of Coyotes, Canis latrans, threatens a remnant herd of Caribou, Rangifer tarandus, in southeastern Québec. Canadian Field-Naturalist 109: 227-235. Georges, S. 1976. A range extension of the Coyote in Québec. Canadian Field-Naturalist 90: 78-79. Lariviére, S., and M. Créte. 1993. The size of eastern Coyotes (Canis latrans): a comment. Journal of Mam- malogy 74: 1072-1074. Lehman, N., A. Eisenhawer, K. Hansen, L. D. Mech, R. O. Peterson, P. J. P. Gogan, and R. K. Wayne. 1991. Introgression of coyote mitochondrial DNA into sympatric North American Grey Wolf populations. Evolution 45: 104-119. Nowak, R. M. 1979. North American Quaternary Canis. Monograph of the Museum of Natural History, Univer- sity Arkansas, number 6, Arkansas. Parker, G. R. 1995. Eastern Coyote: The story of its success. Nimbus Publishing Ltd., Halifax, Nova Scotia. 256 pages. Parker, G. R., and S. Luttich. 1986. Characteristics of the Wolf (Canis lupus labradorius Goldman) in Northern Quebec and Labrador. Arctic 39: 145-149. Patterson, B. R., S. B.-Nielsen, and F. Messier. 1999. Activity patterns and daily movements of the Eastern NOTES 129 Coyote, Canis latrans, in Nova Scotia. Canadian Field- Naturalist 113: 251-257. Patterson, B. R., L. K. Benjamin, and F. Messier. 2000. Winter nutritional condition of eastern coyotes in rela- tion to prey density. Canadian Journal of Zoology 78: 420-427. Perkins, P. J., and W. W. Mautz. 1990. Energy require- ments of eastern coyotes. Canadian Journal of Zoology 68: 656-659. Pilgrim, K. L., D. K. Boyd, and S. H. Forbes. 1998. Testing for wolf-coyote hybridisation in the Rocky Mountains using mitochondrial DNA. Journal of Wild- life Management 62: 683-689. Poulle, M.-L., M. Créte, and J. Huot. 1995. Seasonal variation in body mass and composition of eastern coy- otes. Canadian Journal of Zoology 73: 1625-1633. Rand, A. L. 1945. Mammals of the Ottawa district. Canadian Field-Naturalist 59: 111-132. Thurber, J. M., and R. O. Peterson. 1991. Changes in body size associated with range expansion in the Coyote (Canis latrans). Journal of Mammalogy 72: 750-755. Tremblay, J. P., M. Créte, and J. Huot. 1998. Summer foraging behaviour of eastern coyotes in rural versus for- est landscape: A possible mechanism of source-sink dynamics. Ecoscience 5: 172-182. Received 24 February 2000 Accepted 25 February 2002 Long Distance Movement by a Coyote, Canis latrans, and Red Fox, Vulpes vulpes, in Ontario: Implications for Disease-spread RICHARD C. ROSATTE Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, Science Complex. P.O. Box 4840, Peterborough, Ontario, K9J 8N8 Canada Rosatte, Richard C. 2002. Long distance movement by a Coyote, Canis latrans, and Red Fox, Vulpes vulpes, in Ontario: Implications for disease-spread. Canadian Field-Naturalist 116(1): 129-131. During a rabies control program in southern Ontario, Raccoons, (Procyon lotor lotor), Striped Skunks (Mephitis mephitis), Red Foxes (Vulpes vulpes), and Coyotes (Canis latrans) were live-captured, vaccinated, ear-tagged, and released at point of capture. One of eight Coyotes captured and released during 1995 in Niagara Falls, Ontario, dispersed 320 km to Coatsworth, Ontario. Additionally, 1 of 23 foxes, captured and released in Scarborough, Ontario, during 1994, moved 170 km to Rossmore, Ontario. Although such long distance movements are probably rare in Ontario they may play a critical role in the dissemination of infectious diseases such as rabies. Key Words: Red Fox, Vulpes vulpes, Coyote, Canis latrans, rabies, movement, dispersal, Ontario. As part of a Provincial strategy to prevent raccoon rabies from becoming established in Ontario, the Ontario Ministry of Natural Resources (OMNR) implemented a Trap-Vaccinate-Release (TVR) pro- gram in the Niagara Frontier (Rosatte et al. 1997). Since 1994, personnel from the OMNR Rabies Research Unit have been live-capturing and vacci- nating Raccoons (Procyon lotor), Striped Skunks (Mephitis mephitis), Red Foxes (Vulpes vulpes) and Coyotes (Canis latrans) in a 680 km? area encom- passing Niagara Falls, Ontario. During the 1995 Niagara TVR program, eight Coyotes were live- captured, vaccinated, ear-tagged, and released at the point of capture in the City of Niagara Falls. One of the Coyotes, a juvenile male, released on 26 July 1995, was shot by a hunter near the town of Coats- worth, Ontario, during February 1996; a straight line distance of 320 km west (Figure 1). All local 130 THE CANADIAN FIELD-NATURALIST. Vol. 116 WE pears % Sock NEW YORK STATE FiGuRE 1. Dispersal direction of a Coyote and a Red Fox in Southern Ontario. humane societies and animal control agencies in the Niagara TVR area were questioned and none had relocated any Coyotes between July 1995 and February 1996. From 1987-1997, the OMNR Rabies Research Unit conducted a TVR program to control rabies in Skunks in the city of Scarborough, Ontario. About 3200 Striped Skunks, 11 300 Raccoons and 100 Red Foxes were live-captured, vaccinated, ear-tagged, and released (Rosatte et al. 1992). During the 1994 Scarborough TVR program, three Red Foxes were captured, vaccinated, ear-tagged, and released at point of capture. On 2 January 1996, a trapper cap- tured one of the foxes near Rossmore, Ontario; a straight line distance of 170 km east (Figure 1). This fox, an adult female in 1996, was originally captured as a juvenile in Scarborough on 19 August 1994. During 1996-1999, 127 Red Foxes and 13 Coyotes were ear-tagged during TVR operations conducted by OMNR in Niagara, Scarborough, and Brockville. No significant movements such as those noted above were reported. Discussion In Ontario, juvenile Red Foxes and Coyotes usual- ly disperse during their first fall or winter (Voigt 1987; MacDonald and Voigt 1985; Voigt and Berg 1987). Researchers in one rural Ontario study deter- mined that Coyote movements ranged from 16 km to 152 km (Kolenosky et al. 1978). Dispersal of juve- nile male and female Red Foxes in another study in rural Ontario averaged 30 km and 8 km, respective- ly. Few individuals dispersed more than 100 km (one juvenile male Red Fox dispersed 122 km with > 120 total Foxes collared) (Voigt 1987; Voigt and MacDonald 1984). Similarly, in Toronto, long dis- tance dispersal was rare; only 2 of 33 radio-collared Red Foxes dispersed over 100 km (Rosatte unpub- lished; Rosatte et al. 1991). Although the movements noted in this study (320 km for the Coyote and 170 km for the Red Fox) are exceptional for Ontario, similar movements have been reported elsewhere. For example, Carbyn and Paquet (1986) recorded a 544 km movement of an adult female Coyote that dispersed from Manitoba to 2002 Saskatchewan. Additionally, one sub-adult male Red Fox moved 395 km from Wisconsin to Indiana (Ables 1965). However, such long distance disper- sals appear to be rare. The last major wildlife rabies epizootic in Canada occurred in the Arctic during the 1940s (Tabel et al. 1974). Numerous wildlife species including Red Foxes and Coyotes were reported with rabies as the disease dispersed into the Canadian provinces from the Northwest Territories in the 1950s (Tabel et al. 1974). The epizootic reached southern Ontario in 1956 due mainly to the movements of infected Red Foxes (Johnston and Beauregard 1969). Fox rabies remained enzootic in southern Ontario since the 1950s, however, the prevalence of that disease has recently declined dramatically due to rabies manage- ment programs involving the use of oral rabies vac- cine baits and TVR (MaclInnes 1987; Rosatte et al. 1992; Rosatte et al. 1993). Dispersing Red Foxes and to a lesser extent Coyotes (as Red Foxes are the pri- mary vector of the Arctic fox strain of rabies) in Ontario have likely played a major role in the main- tenance of the disease as well as in the initiation of new outbreaks (MacInnes 1987; Johnston and Beauregard 1969; Tabel et al. 1974). To minimize the risk of dispersing Red Foxes contributing to the spread of rabies in Ontario, OMNR is annually dis- tributing oral rabies vaccine baits over areas that include a 50 km buffer around any rabies case. Based on observations from this study, long-distance movements by Red Foxes and Coyotes likely are rare. However, biologists should consider the poten- tial maximum dispersal of Red Foxes and Coyotes when planning or implementing a rabies manage- ment strategy for Ontario or other areas where either of these two species occur. Acknowledgments Special thanks to Garnie Skinkle and Richard Provost for reporting the ear-tagged Red Fox and Coyote. The co-operation of the hunters and trap- pers of Ontario is critical to the evaluation of the OMNR rabies program. C. Davies, K. Abraham, M. Obbard, A. Jano, C.D. MacInnes and D. R. Voigt reviewed the manuscript. This is OMNR Wildlife Research and Development Section contribution number 20. Literature Cited Ables, E. D. 1965. An exceptional fox movement. Journal of Mammalogy 46: 102. NOTES 131 Carbyn, L., and P. Paquet. 1986. Long distance move- ment of a coyote from Riding Mountain National Park. Journal of Wildlife Management 50: 89. Johnston, D., and M. Beauregard. 1969. Rabies epi- demiology in Ontario. Bulletin of the Wildlife Disease Association 5: 357-370. Kolenosky, G. B., D. R. Voigt, and R. O. Standfield. 1978. Wolves and Coyotes in Ontario. Ontario Ministry of Natural Resources publication 18 pages. MacDonald, D. W., and D. R. Voigt. 1985. The biologi- cal basis of rabies models. Pages 71-108. in Population dynamics of rabies in wildlife. Edited by P. Bacon. Academic Press publishers. Toronto. MaclInnes, C.D. 1987. Rabies. Pages 910-929 in Wild Furbearer Management and Conservation in North America. Edited by M. Novak, J. Baker, M. Obbard, and B. Malloch. Ontario Trappers Association Publishers, North Bay, Ontario. Rosatte, R. C., M. J. Power, and C. D. MacInnes. 1991. Ecology of urban skunks, foxes and raccoons in Metropolitan Toronto. Pages 31-38 in Wildlife Conser- vation in Metropolitan Environments. Edited by L. W. Adams and D.L. Leedy. National Institute for Urban Wildlife Publishers, Columbia, Maryland. Rosatte, R. C., M. J. Power, C. D. MacInnes, and J. B. Campbell. 1992. Trap-vaccinate-Release and oral vac- cination for rabies control in urban skunks, raccoons and foxes. Journal of Wildlife Diseases 28: 562-571. Rosatte, R. C., C. D. MacInnes, M. J. Power, D. H. Johnston, P. Bachmann, C. P. Nunan, C. Wannop, M. Pedde, and L. Calder. 1993. Tactics for the control of wildlife rabies in Ontario Canada. Scientific and Technical Reviews of the Office of International Epi- zootics 12(1): 95—98. Rosatte, R. C., C. D. MacInnes, R. Taylor Williams, and O. Williams. 1997. A proactive prevention strategy for raccoon rabies in Ontario, Canada. Wildlife Society Bulletin 25: 110-116. Tabel, H., H. Corner, W. Webster, and C. Casey. 1974. History and epizootiology of rabies in Canada. Canadian Veterinary Journal 15: 271-281. Voigt, D. R. 1987. Red Fox. Pages 379-392 in Wild Furbearer Management and Conservation in North America. Edited by M. Novak, J. Baker, M. Obbard, and B. Malloch. Ontario Trappers Association Publishers, North Bay, Ontario. Voigt, D. R., and D. W. MacDonald. 1984. Variation in the spatial and social behaviour of the red fox, Vulpes vulpes). Acta Zoologica Fennica 171: 261-265. Voigt, D. R., and W. E. Berg. 1987. Coyote. Pages 345-357 in Wild Furbearer Management and Conser- vation in North America. Edited by M. Novak, J. Baker, M. Obbard, and B. Malloch. Ontario Trappers Associ- ation Publishers, North Bay, Ontario. Received 14 August 2000 Accepted 20 March 2002 132 THE CANADIAN FIELD-NATURALIST. Vol. 116 Predation by Wolves, Canis lupus, on Wolverines, Gulo gulo, and an American Marten, Martes americana, in Alaska. KEVIN S. WHITE!:4, HoOwARD N. GOLDEN!, Kris J. HUNDERTMARK2, and GERALD R. LEE? !Alaska Department of Fish and Game, Division of Wildlife Conservation, 333 Raspberry Road, Anchorage, Alaska 99518 USA 2Alaska Department of Fish and Game, Division of Wildlife Conservation, 43961 Kalifornsky Beach Road, Suite B, Soldotna, Alaska 43961 USA 3Basin Airmotive, P.O. Box 148, Glennallen, Alaska 99588 4Present address: Alaska Department of Fish and Game, Division of Wildlife Conservation, PO Box 240020, Douglas, Alaska 99824, U.S.A. (e-mail:Kevin_white @ fishgame.state.ak.us) White, Kevin S., Howard N. Golden, Kris J. Hundertmark, and Gerald R. Lee. 2002. Predation by Wolves, Canis lupus, on Wolverines, Gulo gulo, and an American Marten, Martes americana, in Alaska. Canadian Field-Naturalist 116(1): 132-134. We report three instances of Wolf predation on mustelids in Alaska; two involved Wolverines and another involved an American Marten. Such observations are rare and in previous studies usually have been documented indirectly. This account provides insight into the potential role of: Wolves in influencing mesocarnivore communities in northern environ- ments. Key Words: Wolf, Canis lupus, Wolverine, Gulo gulo, Marten, Martes americana, predation, Alaska. Observations of Wolf (Canus lupus) -carnivore inter- actions are rare and have generally focused on those concerning ursids (Ursus arctos, Grizzly Bear; and U. americana, Black Bear; Murie 1944; Mech 1970, Rogers and Mech 1981; Ballard 1982) and canids (Canis latrans, Coyotes, and Vulpes vulpes, Red Foxes; Stenlund 1955; Mech 1970; Berg and Chesness 1978; Peterson 1996; Crabtree and Sheldon 1999). Nevertheless, direct and indirect evi- dence suggests that interactions involving actual and attempted Wolf predation on mustelids (Stenlund 1955, Boles 1977, Route and Peterson 1991, Boyd et al. 1994, Paragi et al. 1996, Kohira and Rextad 1997; Mech et al. 1998) can occur at low frequency in some regions of North America and, further, may have a significant effect on mustelid populations (Palomares and Caro 1999). We report here on observations of Wolverine, Gulo gulo, and American Marten, Martes americana, predation by Wolves in Alaska. Account 1: On 20 March 1983 (1330 PST), during an aerial sex and age composition survey for Moose (Alces alces), we observed an adult Wolverine in the top of a large cottonwood (Populus trichocarpa) tree along the Chilkat River near Haines in southeastern Alaska (59° 37'N, 135° 55'W). Near the base of the tree was a pack of at least five Wolves in an area of blood-stained snow where the Wolves appeared to have been digging. One Wolf was observed with a juvenile Wolverine in its mouth. A few of the other . Wolves were digging in the snow at what we sus- pected was a Wolverine den site, while another Wolf was lying down a short distance from the activity. Our impression was that the Wolves found the Wolverine den by chance and were in the process of digging it out to get to the kits. We were unable to determine if the Wolves actually consumed any of the kits, but judging from the extent and dispersion of blood on the snow, more than one kit had been killed. Account 2: We believe Wolves were responsible for the death of a yearling female Wolverine in an area of light spruce (Picea spp.) forest and tundra vegeta- tion in the Nelchina Basin in southcentral Alaska (62° 41'N, 147° 45'W). On 1 June 1997, while radio- tracking Wolverines as part of a population ecology study (Golden 1997), we observed a Wolf circling a yearling female Wolverine whose radio collar was on mortality mode and which showed no sign of movement (S. D. Bowen, Alaska Department of Fish and Game, personal communication). This was the first day we had detected her radio signal on mortali- ty mode since she was last seen alive on 15 May 1997. We retrieved the carcass by helicopter on 2 June 1997. While the carcass was being loaded onto the helicopter, two Wolves stood within 300 m bark- ing and howling (J. W. Testa, Alaska Department of Fish and Game, personal communication). The necropsy revealed five puncture holes in the skin, three in the chest and two in the groin, which may have been made by canine teeth of Wolves. Its chest was crushed laterally on the ventral side, resulting in several broken ribs. Although the carcass was in an advanced state of decomposition, it was intact and no part of it had been consumed. These observations plus the behavior of the Wolves and the timing of the death in late May suggest Wolves attacked and killed the Wolverine, possibly in defense of a wolf den site. Account 3: On 8 June 2000 (1222 ADT) during a telemetry re-location flight, we observed a radio- 2002 collared, solitary female Wolf in mixed-spruce (P. glauca and P. mariana) forest vigorously digging in moss-covered, hummocky soil, near Old Man Lake, Alaska (62° 31'N, 146° 81'W). As we circled the Wolf, we noticed one American Marten escaping through the forest ~15 m away as the Wolf contin- ued digging in the original location. Subsequently, after ~2 minutes of digging, we observed the Wolf drag another Marten with its jaws from the under- ground cavity that it had been excavating. The Wolf then repeatedly bit, shook and dropped the Marten 4-5 times until the Marten stopped moving, at which time it was presumed dead. The Wolf first stood guarding, and then rolled on the carcass until we ter- minated our observation at 1231 (ADT). Our observations indicated, to the extent possible, that Wolves did not feed on the carcasses of the Wolverines they had killed, a behavior noted by oth- ers (Burkholder 1962, Boles 1977). We can only speculate on the basis of that behavior, but ultimate explanations might include: elimination of competi- tors, defense of offspring, availability of prey, or dis- turbance by human observers. The role of Wolves in structuring mesocarnivore communities is suspected to be significant though not fully understood. In areas where Wolves are re- colonizing historic ranges following prolonged absences, behavioral and ecological modifications of mesocarnivores can be dramatic (Crabtree and Sheldon 1999). Such changes underscore the impor- tance of Wolves in influencing ecosystem dynamics in such areas. While Wolf populations have fluctuat- ed historically in Alaska, their presence has remained constant and their role in influencing ungu- late populations has been studied extensively (Gasaway et al. 1983; Ballard et al. 1992; Dale et al. 1994; Adams et al. 1995), though their interactions with mesocarnivores has received little attention. When considering species such as Wolverines that typically occur at low density, it is important to rec- ognize the role that even limited predation might exert on their population dynamics. Thus, in such cases, anecdotal accounts of predator-specific mor- tality provide valuable insights into the range of influence that Wolves might exert on mesocarnivore communities in relatively undisturbed northern envi- ronments. Acknowledgments We thank S.D. Bowen, J. W. Testa, and D. Mortenson for their assistance with field observa- tions. We also thank S. Pyare and two anonymous reviewers for providing insightful comments on this manuscript. This work was funded by the Alaska Department of Fish and Game and Federal Aid to Wildlife Restoration Grant W-24-3, W-24-4, W-24- 5, and W-27-1. NOTES 133 Literature Cited Adams, L. G., B. W. Dale, and L. D. Mech. 1995. Wolf predation on caribou calves in Denali National Park, Alaska. Pages 245-260 in Ecology and conservation of wolves in a changing world. Edited by L. N. Carbyn, S.H. Fritts, and D. R. Seip. Canadian Circumpolar Institute, Occasional Publication Number 35. Ballard, W. B. 1982. Gray wolf-brown bear relationships in the Nelchina Basin of south-central Alaska. Pages 71-80 in Wolves of the world: perspectives of behavior, ecology and conservation. Edited by F. H. Harrington, and P. C. Paquet. Noyes Publications, Park Ridge, New Jersey, USA. Ballard, W. B., J. S. Whitman, and D. J. Reed. 1991. Population dynamics of moose in south-central Alaska. Wildlife Monographs 114. Berg, W. E., and R. A. Chesness. 1978. Ecology of coy- otes in northern Minnesota. Pages 229-247 in Coyotes: biology, behavior, and management. Edited by M. Beckoff. Academic Press, New York, USA. Boles, B. K. 1977. Predation by wolves on wolverines. Canadian Field-Naturalist 91: 68-69. Boyd, D. K., R. R. Ream, D. H. Pletcher, and M. W. Fairchild. 1994. Prey taken by colonizing wolves and hunters in the Glacier National Park area. Journal of Wildlife Management 58: 289-295. Burkholder, B. L. 1962. Observations concerning wol- verine. Journal of Mammalogy 43: 263-264. Crabtree, R. L., and J. W. Sheldon. 1999. Coyotes and canid coexistence in Yellowstone. Pages 127-163 in Carnivores in ecosystems: the Yellowstone experience. Edited by T. W. Clark, A. P. Curlee, S.C. Minta, and P.M. Kareiva. Yale University Press, New Haven, Connecticut, USA. Dale, B. W., L. G. Adams, and R. T. Bowyer. 1994. Functional response of wolves preying on barren-ground caribou in a multiple-prey ecosystem. Journal of Animal Ecology 63: 644-652. Gasaway, W. C., R. O. Stephenson, J. L. Davis, P. E. K. Shepherd, and O. E. Burris. 1983. Interrelationships of wolves, prey, and man in interior Alaska. Wildlife Monographs 84. Golden, H. N. 1997. Furbearer management techniques development: densities, trend, and harvest potential of wolverine populations. Alaska Department of Fish and Game, Federal Aid to Wildlife Restoration Research Grant W-24-5, Research Progress Report. Kohira M., and E. A. Rexstad. 1997. Diets of wolves, Canis lupus, in logged and unlogged forests of south- eastern Alaska. Canadian Field-Naturalist 111: 429-435. Mech, L. D. 1970. The wolf: ecology and behavior of an endangered species. Doubleday, New York, USA. Mech, L. D., L. G. Adams, T. J. Meier, J. W. Burch, and B. W. Dale. 1998. The wolves of Denali. University of Minnesota Press. Minneapolis, USA. Murie, A. 1944. The wolves of Mount McKinley. U.S. National Park Service, Fauna Series Number 5. Palomares, F., and T. M. Caro. 1999. Interspecific killing among mammalian carnivores. American Naturalist 153: 492-508. Paragi, T. F., W. N. Johnson, D. D. Katnik, and A. J. Magoun. 1996. Marten selection of postfire seres in the Alaskan taiga. Canadian Journal of Zoology 74: 2226-2237. 134 THE CANADIAN FIELD-NATURALIST Peterson, R. O. 1996. Wolves as interspecific competi- tors in canid ecology. Pages 315-323 in Ecology and conservation of wolves in a changing world. Edited by L. N. Carbyn, S.H. Fritts, and D. R. Seip. Canadian Circumpolar Institute, Occasional Publication Number a5. Rogers L. L., and L. D. Mech. 1981. Interactions of wolves and black bears in northeastern Minnesota. Journal of Mammalogy 62: 424-436. Route, W. T., and R. O. Peterson. 1991. An incident of Vol. 116 wolf, Canis lupus, predation on a River Otter, Lutra canadensis, in Minnesota. Canadian Field-Naturalist 105: 567-568. Stenlund, M. H. 1955. A field study of the timber wolf (Canis lupus) on the Superior National Forest, Min- nesota. Minnesota Department of Conservation, Tech- nical Bulletin Number. 4. Received 1 August 2000 Accepted 25 March 2002 Meek’s Halfbeak, Hyporhamphus meeki, and Flying Gurnard, Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia A. JAMIE F. Gipson! and RANSOM A. MYERS? \Acadia Centre for Estuarine Research, P.O. Box 115, Acadia University, Wolfville, Nova Scotia BOP 1X0 Canada; e-mail: jamie.gibson @acadiau.ca [Author to whom correspondence should be addressed] "Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 3J5 Canada; e-mail: myers@mscs.dal.ca Gibson, A. Jamie F., and Ransom A. Myers. 2002. Meek’s Halfbeak, Hyporhamphus meeki, and Flying Gurnard Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia. Canadian Field-Naturalist 116(1): 134-135. We report the capture of several unusual fish at a hydroelectric generating station, Annapolis Royal, in the Annapolis Basin, Nova Scotia, during the fall of 1999. These include a Meek’s Halfbeak, Hyporhamphus meeki, that is either the first or second Canadian record for this species and a Flying Gurnard, Dactylopterus volitans, that is a first record for the Bay of Fundy. Additionally, three Bluefish, Pomatomus saltatrix, and a Fourbeard Rockling, Enchelyopus cimbrius, were reported for the first tume from the Annapolis Estuary during 1999. Key Words: Meek’s Halfbeak, Hyporhamphus meeki, Flying Gurnard, Dactylopterus volitans, Bluefish, Pomatomus salta- trix, Fourbeard Rockling, Enchelyopus cimbrius, Annapolis River, Bay of Fundy, Nova Scotia, distribution. The Annapolis Basin in Nova Scotia is home to the western hemisphere’s only tidal hydroelectric gener- ating station, located in Annapolis Royal (44° 45’N, 65° 31’W). As such, fish assemblages within the estuary are well studied, during both pre-operational stock assessments and surveys (e.g., Melvin et a. 1985; Jessop 1976), and several assessments of fish passage facilities at the generating station (e.g., Gibson 1996). During 1999, several unusual fish specimens were captured while testing the effective- ness of an ultrasound fish diversion system at the sta- tion (Gibson and Myers 2000). A Meek’s Halfbeak, Hyporhamphus meeki, captured on 23 September, 1999, is either the first or second record of this species in Canada, and is the first record in 50 years. A Flying Gurnard, Dactylopterus volitans, captured 28 September 1999, is a first record for the Bay of Fundy. Other unusual specimens captured at this location during September 1999 include three. Bluefish, Pomatomus saltatrix and one Fourbeard rockling, Enchelyopus cimbrius. While these latter species occur regularly in the Bay of Fundy (Scott and Scott 1988), these species have not been reported previously in the Annapolis Estuary. Hyporhamphus spp. are planktivorous fish charac- terised by a very long lower jaw and short upper jaw. The taxonomic status of western Atlantic Hyporham- pus was clarified by the description of a new species, Hyporhamphus meeki, by Banford and Collette (1993). This species ranges north from the Gulf of Mexico, and is a rare stray into the Gulf of Maine. It usually can be distinguished from its southern rela- tive, Hyporhamphus unifasciatus, by having a greater number of total gill rakers on the first (31 to 40) and second arches (20 to 30), and having a ratio of preor- bital length to orbit diameter greater than 0.70 (Banford and Collette 1993). Contreras-Balderas et al. (1997) suggest the species can be distinguished using the shape of the lateral band. The specimen reported herein is unusual in having 29 gill rakers on right first arch and 23 gill rakers on the right second arch, both of which are low for this species. The identifying characteristic for this specimen is the ratio of preor- bital length to orbit diameter, which equals to 0.74 (Bruce Collette, personal communication). One previous record of Hyporhamphus sp. exists for eastern Canada. This specimen, reported as Hyporhamphus unifasciatus, was captured in a 2002 herring weir at Chamcook, New Brunswick, on 30 September, 1949 (Leim and Day 1959). Its location suggests that this specimen may actually have been a H. meeki, a species that was not described at the time that this specimen was captured. Dactylopterus volitans is a benthic species of fish that superficially resembles a searobin, and is typically found in tropical or warm temperature water. It ranges south to Argentina, and commonly is found as far north as North Carolina. The species occasionally strays north to Nova Scotia. Scott and Scott (1988) report four specimens from the Atlantic coast of Nova Scotia during the 1970s. The Nova Scotia Museum of Natural History has two records of D. volitans in Nova Scotia, both from the Atlantic Coast in 1990 (John Gilhen, personal communication). D. volitans have also been reported from the Grand Banks and Scotian Shelf (Lou VanGuelpen, personal communication). Bluefish are regular summer visitors to the Bay of Fundy during warmer summers. The Fourbeard Rockling is a resident of the Bay of Fundy, and some- times moves into embayments during summer and fall (Scott and Scott 1988). Water temperature was warmer and salinity was higher during early September in 1999 than during six previous years of sampling since 1985, perhaps leading to favourable conditions for these fish to move farther into the estuary. The Meek’s Halfbeak (reference number: ARC 0016423) and Flying Gurnard (reference number: ARC 0016422) are deposited at the Atlantic Reference Centre located at the Huntsman Marine Science Centre, St. Andrews, New Brunswick. Acknowledgments The authors express their thanks to John Gilhen (Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia, B3H 3A6), Lou VanGuelpen (Atlantic Reference Centre, Huntsman Marine Science Centre, 1 Lower Campus Road, St. Andrews, New Brunswick, E5B 2L7), Donald F. McAlpine (New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick, E2K 1E5) for searching their collections for records of these species. Thanks are also extended to Bruce Collette (Systematics Laboratory, National Marine Fisheries Services, Smithonian Institution, NOTES 135 Washington, DC, 20560) for assistance with the identification of the H. meeki. Literature Cited Banford, H. M., and B. B. Collete. 1993. Hyporhamphus meeki, A new species of halfbeak (Teleostei: Hemiramphidae) from the Atlantic and Gulf coasts of the United States. Proceedings of the Biological Society of Washington 106: 369-384. Contreras-Balderas, S., M. L. Lozano-Vilano, and M. E. Garcia Ramirez. 1997. Distributional and ecological notes on the halfbeaks of eastern Gulf of Mexico, with a provisional key for their identification. Gulf Research Reports 9: 327-331. Gibson, A. J. F. 1996. Distribution and seaward migra- tion of young-of-the-year American Shad (Alosa sapidissima), Blueback Herring (A. aestivalis) and Alewives (A. psuedoharengus) in the Annapolis River Estuary. M. Sc. thesis. Acadia University, Wolfville, Nova Scotia. 96 pages. Gibson, A. J. F., and R. A. Myers. 2000. Assessment of a high-frequency sound fish diversion system at the Annapolis Tidal Generating Station with notes about the survival of fish moving seaward at the Annapolis cause- way. Acadia Centre for Estuarine Research Publication Number 55. Acadia University, Wolfville, Nova Scotia. 98 pages. Jessop, B. M. 1976. Physical and Biological Survey of the Annapolis River, 1975. Data Record Series Number MAR/D-76-8. Fisheries and Marine Service, Department of the Environment, Halifax. 29 pages. Leim, A. H., and L. R. Day. 1959. Records of uncommon and unusual fishes from eastern Canadian waters. 1950- 1958. Journal of the Fisheries Research Board of Canada 16: 503-514. Melvin, G. D., M. J. Dadswell, and J. D. Martin. 1985. Impact of Lowhead Hydroelectric Tidal Power Development on Fisheries I. A Pre-operation Study of the Spawning Population of American Shad Alosa sapidissima (Pisces:Clupeidae). in the Annapolis River, Nova Scotia, Canada. Canadian Technical Report of Fisheries and Aquatic Sciences Number 1340. St. Andrews, New Brunswick. 33 pages. Scott, W. B., and M. G. Scott. 1988. Atlantic Fishes of Canada. Canadian Bulletin of Fisheries and Aquatic Sciences 219. 731 pages. Received 5 June 2000 Accepted 19 February 2002 136 THE CANADIAN FIELD-NATURALIST Vol. 116 Apparent Capture Myopathy in Hoary Bats, Lasiurus cinereus: A Cautionary Note Tuomas S. JuNG!2, IAN D. THompson3, M. BRIAN C. HICKEY* and RODGER D. TITMAN! \McGill University, Department of Natural Resource Sciences, 21111 Lakeshore Blvd., Montreal, Quebec, H9X 3V9 Canada 2Present address: Yukon Department of Renewable Resources, Fish and Wildlife Branch, P.O. Box 2703, Whitehorse, Yukon, Y1A 2C6 Canada; thomas.jung @ gov.yk.ca 3Canadian Forest Service, Great Lakes Forestry Centre, 1219 Queen Street East, Sault Ste. Marie, Ontario, P6A 5M7 Canada 4Y ork University, Department of Biology, 4700 Keele St., North York, Ontario, M3J 1P3 Canada 5Present address: St. Lawrence River Institute of Environmental Sciences, Cornwall, Ontario, KOH 1E1 Canada Jung, Thomas S., Ian D. Thompson, M. Brian C. Hickey, and Roger D. Titman. 2002. Apparent capture myopathy in Hoary Bats, Lasiurus cinereus: a cautionary note. Canadian Field-Naturalist 116(1): 136-137. Capture myopathy, a stress-induced disease resulting in metabolic acidosis, has not been recorded in bats. We observed two cases of apparent capture myopathy in Hoary Bats (Lasiurus cinereus) captured in mist-nets. We caution those intend- ing to work with this species of a potential occurrence, and discuss capture protocols that may reduce the risk of capture myopathy. Key Words: Hoary Bats, Lasiurus cinereus, capture myopathy, live-capture. Biologists often rely on data obtained through live- capture. Live-capturing animals, however, may result in capture myopathy: a stress-induced disease, where- by the liberation of large amounts of lactic acid results in metabolic acidosis, which may lead to mus- cle necrosis and death (Chalmers and Barrett 1982; Pond and O’Gara 1994). Clinical signs of this condi- tion may include muscle stiffness, loss of coordina- tion, listlessness, and increased respiration and body temperatures, but affected animals can die without exhibiting outward symptoms (Chalmers and Barrett 1982). Capture myopathy can occur immediately upon capture or up to one month afterwards, and has been reported from a variety of species. To the best of our knowledge, there are no published reports of bats succumbing to capture myopathy. We encourage others to collate and disseminate their observations of capture myopathy in bats. As part of a study on the habitat ecology of forest- dwelling bats in central Ontario (46.8° N; 81.9° W; Jung et al. 1999), bats were live-captured in mist- nets set across rivers and logging roads. On 4 August 1995 at 22:40 (EDT) an adult male Hoary Bat was captured. The bat was extracted from the net in <5 min, and was active and vocal when captured. Once extracted from the net, the bat was placed in a home- made steel mesh holding container (Kunz and Kurta 1988), for about 25 min. After which the bat was dis- covered languid and did not fly. Therefore, it was placed on the limb of a small Trembling Aspen (Populus tremuloides), where it remained, suspended by its hind feet. The next morning the bat was dis- covered dead. Post-mortem examination indicated that the bat was in good condition, as the pelage was shiny, there were few noticeable ectoparasites, and its mass appeared normal (29.8 g.). Due to the diffi- culty in keeping the specimen frozen at a remote field camp, it was not sent for necropsy. We do not believe that the Hoary Bat was excessively handled, relative to > 100 Myotis spp. that we had captured. It is our belief that the Hoary Bat died as a result of the stress of being captured. Similarly, as part of a study of the thermal ecology and foraging behavior of female Hoary Bats in southwestern Ontario (43.3° N; 81.8° W; Hickey and Fenton 1996), Hoary Bats were live-captured near street lights by swinging mist-nets at diving bats. On 8 June 1989, an adult female Hoary Bat was cap- tured and banded. After about 10 min the bat was hand-released, but was listless and did not fly away. The bat was placed on the limb of a nearby tree and it remained there. The bat was checked daily and had not moved. On 12 June 1989, the bat was discovered dead, having apparently not moved since being cap- tured. The bat did not appear injured, and seemed to have perished as a result of capture myopathy. In a related manner, our experience (MBCH) with Hoary Bats in captivity suggests that these species have a propensity to die within 1 month of capture, which may also be stress-related. This is the first known report of apparent stress- induced capture myopathy in bats. We do not know why Hoary Bats may be more susceptible to capture myopathy than other bats such as Myotis spp. Although the prognosis is poor for individuals exhibit- ing signs of capture myopathy, Pond and O’Gara (1994) state that keeping the animal well-oxygenated and providing intravenous or subcutaneous adminis- tration of sodium bicarbonate in a saline solution is a recommended therapy. If stress-induced capture myopathy does occur somewhat regularly in Hoary Bats, then the use of a chemical immobilization agent 2002 NOTES 37 may reduce the likelihood of capture myopathy, through a reduction in the onset of extreme stress. Chemical tranquilizers have been used for small mammals (e.g., Wright 1983), but are not commonly used on bats. We recommend that Hoary Bats be pro- cessed as quickly as possible after live-capture, as they may not be as resilient to capture as other species of bats. We also suggest that steel mesh containers may provide substantially more oxygen than cloth bags, which are sometimes used to hold bats, thereby reducing the likelihood of capture myopathy. Bats that do perish after capture should be sent for necropsy to verify the cause of death. Acknowledgments R. M. Brigham provided thoughtful comments on an earlier draft of this note. We thank the field assis- tants that participated in mist-netting. Live-capture of bats was conducted under the approval of animal care committees at McGill and York universities. Literature Cited Chalmers, G. A., and M. W. Barrett. 1982. Capture myopathy. Pages 84—94. Jn Noninfectious diseases of wildlife. Edited by G. L. Hoff and J. W. Davis. Iowa State University Press. Ames, Iowa. 174 pages. Hickey, M. B. C., and M. B. Fenton. 1996. Behavioural and thermoregulatory responses of female Hoary Bats, Lasiurus cinereus, to variations in prey availability. Ecoscience 3: 412-422. Jung, T.S., I. D. Thompson, R. D. Titman, and A. P. Applejohn. 1999. Habitat selection by forest-dwelling bats in relation to stand type and structure in central Ontario. Journal of Wildlife Management 63: 1306-1319. Kunz, T. H., and A. Kurta. 1988. Capture methods and holding devices. Pages 1-30 In Ecological and behav- ioral methods for the study of bats. Edited by T. H. Kunz. Smithsonian Institution Press, Washington, D.C. Pond, D. B., and B. W. O’Gara. 1994. Chemical immobi- lization of large mammals. Pages 125-139 Jn Research and management techniques for wildlife and habitats. Fifth ed. Edited by T. A. Bookhout. The Wildlife Society, Bethesda, Maryland. 740 pages. Wright, J. M. 1983. Ketamine hydrochloride as a chemi- cal restraint for selected small mammals. Wildlife Society Bulletin 11: 76-79. Recieved 9 June 2000 Accepted 22 March 2002 Mobbing Black-billed Magpie, Pica hudsonia, Killed by Cooper’s Hawk, Accipiter cooperii GEOFFREY L. HOLROYD Canadian Wildlife Service, Room 200, 4999-98 Avenue, Edmonton, Alberta T6B 2X3 Canada Holroyd, Geoffrey L. 2002. Mobbing Black-billed Magpie, Pica pica, killed by Cooper’s Hawk, Accipiter cooperii. Canadian Field-Naturalist 116(1): 137-138. A Cooper’s Hawk killed a Black-billed Magpie that was part of a group mobbing it. This occurrence happened before the breeding season. Any benefit of mobbing did not enhance the survival of this magpie. Key Words: Black-billed Magpie, Pica hudsonia, Cooper's Hawk, Accipiter cooperii, mobbing, mortality. Mobbing is the massing together of birds, often of different species, for the purpose of aggression and attack against a common predator (Terres 1982). Mobbing is most prevalent in the nesting season (Shedd 1982) but does occur year-round, particularly by species that remain on their nesting territory (Shedd 1983). This note documents an observation of a Cooper’s Hawk (Accipiter cooperii) killing and eat- ing a Black-billed Magpie (Pica hudsonia) that was one of a group that was mobbing it. On 15 April 2000, just after 17:00 h, Phil Trefry drew our attention to a Cooper’s Hawk on a White Birch (Betula papyrifera) in a low lying birch-willow (Salix spp.) bluff in a depression in front of his home about 50 km east of Edmonton (54° 24'N 112° 50'W). The farm is in rolling aspen parkland with open fields and Trembling Aspen (Populus tremu- loides) and Balsam Poplar (P. balsamifera) dominat- ed woods. The hawk was being mobbed by four mag- pies and two Blue Jays (Cyanocitta cristata). The mobbing was mostly “vocal approach” (sensu Shedd 1982) and flying at a distance. The mobbers did not fly at or strike the hawk. A Red-tailed Hawk (Buteo jamaicensis) was perched 50 m west of the Cooper’s Hawk, higher in a birch tree, but was ignored by the corvids. One agitated magpie perched 1-2 m above the Cooper’s Hawk in the same tree, calling and hop- ping (“mobbing” sensu Shedd 1982). The hawk glid- ed to the ground under overhanging willow branches, 10 m to the south of its original perch. The magpie hopped to the ground about 3 m from the hawk where they were hidden from the observers by tall dead herbaceous vegetation. The hawk flew to the magpie, killed it and carried it along the edge of the bluff, 138 50 m north, then into the bluff behind some birch trees and out of sight. Over 10 magpies and two Blue Jays arrived from different directions and they and the other corvids called, hopped agitatedly and flew over- head. The Red-tailed Hawk and another that had been perched to the east, flew closer and perched nearby. After about 10 minutes the corvids and red-tails left. Three hours later we visited the two sites. At the kill site we could see the wing-tip prints of the Cooper’s Hawk in the snow and two marks between the prints, presumably where the magpie had landed and died. At the northerly destination was a collection of magpie body feathers over an area of 3 sq. m, where the hawk had plucked part of the magpie. There were no flight feathers nor any of the magpie's carcass. The purpose of mobbing is to drive a predator from a nearby nest or territory (Shedd 1982). Mobbing birds have been observed driving aerial predators to the ground or into the water (Terres 1982: 63, 627). However, mobbing does not always drive a‘predator from a nest (Taylor 1972). The present observation is unusual because the magpie was killed while mob- bing and magpies are rarely preyed upon by raptors. In April, Black-billed Magpies can be expected on territory. Although no magpies nested in the bluff where this observation took place, they are common breeding birds in the area (personal observation). Mobbing is more common in the breeding season in migratory species such as the American Robin (Shedd 1982), but occurs year-round with no appar- ent seasonality in non-migratory species such as Black-capped Chickadee (Poecile atricapillus) (Shedd 1983). The presumed advantage of mobbing to a bird without a nest is to drive the potential predator from the permanent territory of the bird. In our observation, the magpie did not have an active nest in the immediate vicinity to protect. Sordahl (1990) reviewed 30 instances of predators killing mobbing birds. At least one of these instances involved a Cooper’s Hawk (Wilson 1986). In 1962 in Virginia, R. D. Denson (1979) observed a Great Horned Owl (Bubo virginianus) kill one of a flock of American Crows (Corvus brachyrhychos) that were mobbing it. The owl grabbed the crow out of the air from its perch. As in our observation, the crow went too close to the owl. Although mobbing behaviour may drive away a predator, the close proximity of predator and potential prey can be expected to have some risk for the prey. In this case the magpie appeared to have made a mistake when it landed on the ground too near the hawk. The magpie was killed with little effort by the Cooper’s Hawk as evidence by the lack of sign of a struggle in the snow. Passerines mobbed Northern Harriers (Circus cya- neus) Only when the harriers are in flight and active- ly hunting (Bildstein 1982). When the harriers are perched, the passerines ignore the hawks. In both our observation and that of Denson (1979) the Great Horned Owl and the Cooper’s Hawk, species that will hunt from a perch, were mobbed while perched. THE CANADIAN FIELD-NATURALIST Vol. 116 The nearby perched red-tails, which are a known nest predator (Trost 1999), were not mobbed. Magpies are rarely taken by raptors (Trost 1999). Three magpies were recorded in 550 prey remains collected at Peregrine Falcon (Falco peregrinus) nests (unpublished data) but none in 203 prey at Prairie Falcon (F. mexicanus) nests in southern Alberta (Hunt 1983). One Cooper’s Hawk nest observed at nearby Beaverhill Lake did not include any magpie remains (unpublished data). Magpies were not reported in the diet of Cooper’s Hawks in Washington and Oregon, which are within the mag- pie's range (Kennedy and Wilson 1986; Reynolds and Meslow 1984). Although this observation is only of a single event, the killing of a mobbing magpie by a Cooper’s Hawk has interesting implications. Acknowledgments I thank Elisabeth Beaubien, Martin Raillard, Tony Erskine and an anonymous reviewer for comments on earlier draft of this manuscript. Elisabeth, Helen Trefry and Phil Trefry also observed this event. Literature Cited Bildstein, K. L. 1982. Responses of Northern Harriers to mobbing passerines. Journal of Field Ornithology 53: 7-14. Denson, R. D. 1979. Owl predation on a mobbing crow. Wilson Bulletin 91: 133. Hunt, L. E. 1983. Diet and habitat use of nesting Prairie Falcons (Falco mexicanus) in an agricultural landscape in southern Alberta. MSc. Thesis, University of Alberta. 60 pages. Kennedy, P. L., and D. R. Wilson. 1986. Prey size selec- tion in nesting male and female Cooper’s Hawks. Wilson Bulletin 98: 110-115. Reynolds, R. T., and E. C. Meslow. 1984. Partitioning of food and niche characteristics of coexisting Accipiter during breeding. Auk 101: 761-779. Shedd, D. H. 1982. Seasonal variation and function of mobbing and related behaviors of the American Robin (Turdus migratorius). Auk 99: 342-346. Shedd, D. H. 1983. Seasonal variation in mobbing inten- sity in the Black-capped Chickadee. Wilson Bulletin 95: 343-348. Sordahl, T. A. 1990. The risks of avian mobbing and dis- traction behaviour: an anecdotal review. Wilson Bulletin 102: 349-352. Taylor, W. K. 1972. Mobbing of a Fish Crow by passe- rines. Wilson Bulletin 84: 98. Terres, J. K. 1982. The Audubon Society Encyclopedia of North American Birds. Knopf, New York. 1109 pages. Trost, C. H. 1999. Black-billed Magpie (Pica pica). In The Birds of North America, Number 389. Edited by A. Poole and F. Gill. The Birds of North America, Inc., Philadelphia, Pennsylvania. 28 pages. Wilson, R. E. 1986. An aerial counter attack by a Cooper’s Hawk. Bulletin of the Texas Ornithological Society 19: 32-33. Received 21 September 2000 Accepted 18 March 2002 2002 NOTES 139 Breeding Season of Wolves, Canis lupus, in Relation to Latitude L. DAvip MECH Biological Resources Division, U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711-37th Street, SE, Jamestown, North Dakota 58401-7317 \Mailing address: The Raptor Centre, University of Minnesota, 1920 Fitch Avenue, St. Paul, Minnesota 55108 USA Mech, L. David. 2002. Breeding season of Wolves, Canis lupus, in relation to latitude. Canadian Field Naturalist 116(1): 139-140. A significant relationship was found between Wolf (Canis lupus) breeding dates and latitudes between 12° and 80° N, with Wolves breeding earlier at lower latitudes, probably because of differences in seasonality. Key Words: Wolf, Canis lupus, reproduction, latitude, breeding, mating. A general relationship between breeding dates in Wolves (Canis lupus) and latitudes from 41°—71°N was noticed by Mech (1970: 117) when he summa- rized published data from several locales. However, he conducted no statistical test of this hypothesis. Herein, I add data from other areas of latitude from 12° to 80° N (Table 1) and statistically test the effect of latitude. I used latitude as the independent variable in a sim- ple linear regression and the reported breeding date as the dependent variable. For breeding date, I used the mid date for the reported breeding season and con- verted all dates to sequential numbers starting with 15 October to facilitate comparing breeding dates before and after the start of the calendar year. The relationship between breeding date and lati- tude was highly significant (r2= 0.74; P < 0.0001; y = 16.19 + 2.23x), supporting Mech’s (1970) hypothesis. On average, breeding season shifts 22 TABLE 1. Mating seasons of wolves at various latitudes. days later with each 10° latitude increase. It seems reasonable to suggest that the shift is related to differ- ences in general seasonality and thus in associated ecological conditions. Acknowledgments This study was supported by the Biological Resources Division of U.S. Geological Survey and the U.S. Department of Agriculture, North Central Research Station. Literature Cited Bailey, V. 1926. A biological survey of North Dakota. U.S. Department of Agriculture, Biological Survey, North American Fauna 49. Cowan, I. M. 1947. The timber wolf in the Rocky Mountain national parks of Canada. Canadian Journal Research 25: 139-174. Fritts, S. H., and L. D. Mech. 1981. Dynamics, move- ments, and feeding ecology of a newly protected wolf Location N Latitude Season Authority Southern India V2" October Kumar and Rahmani 2001 Arizona* 34° February, March W. Brown, personal communication Illinois@ 42° February Rabb 1968 Yellowstone National Park> 45° February Smith et al. 1998 Ontario 47° Early March Joslin 1966 North Dakota 46°—49° January Bailey 1926 Isle Royale (Michigan) 47° Late February Mech 1966 Minnesota 48° February Mech and Knick 1978; Fritts and Mech 1981; Fuller 1989 British Columbia 51°-53° March Cowan 1947 Germany* PA Mid March Schonberner 1965 Alberta 60° February, March Soper 1942; Fuller and Novakowski 1955 Northwest Territories 60°-65° Late March Kelsall 1960 Alaska 60°—71° March Murie 1944; Kelly 1954; Rausch 1967;, Lentfer and Sanders 1973; Mech et al. 1998 Finland 60°-70 March Pulliainen 1965 Russia ai? Late March-early April Makridin 1962 Ellesmere Island 80° Late March-early April Mech 1993 “Captive wolves »Wolves translocated from 53—56° N. 140 THE CANADIAN FIELD-NATURALIST population in northwestern Minnesota. Wildlife Mono- graph 80. Fuller, T. K. 1989. Population dynamics of wolves in north-central Minnesota. Wildlife Monograph 105. Fuller, W. A., and N. S. Novakowski. 1955. Wolf control operations, Wood Buffalo National Park, 1951-1952. Canadian Wildlife Service, Wildlife Management Bul- letin Series 1, Number 11. Joslin, P. W. B. 1966. Summer activities of two timber wolf (Canis lupus) packs in Algonquin Park. M.S. the- sis, University of Toronto, 99 pages. Kelly, M. W. 1954. Observations afield on Alaskan wolves. Proceedings of Alaska Science Conference 5: 35. Kelsall, J. P. 1960. Co-operative studies on barren ground caribou 1957-58. Canadian Wildlife Service, Wildlife Management Bulletin Series 1, Number 12. Kumar, S., and A. R. Rahmani. 2001. Livestock depre- dation by wolves in the great Indian Bustard Sanctuary, Nannaj (Maharashtra), India. Journal, Bombay Natural History Society 97: 340-348. Lentfer, J. W., and D. K. Sanders. 1973. Notes on the captive wolf (Canis lupus) colony, Barrow, Alaska. Canadian Journal of Zoology 51: 623-627. Makridin, V. P. 1962. The wolf in the Yamal north. Zoological Zhurnal 41: 1413-1417. Mech, L. D. 1966. The wolves of Isle Royale. U. S. National Park Service Fauna Series Number 7. Mech, L. D. 1970. The wolf: ecology and behavior of an endangered species. Natural History Press, Doubleday Publishing Co., New York. Vol. 116 Mech, L. D. 1993. Resistance of young wolf pups to inclement weather. Journal of Mammalogy 74: 485-486. Mech, L. D., and S. T. Knick. 1978. Sleeping distances in wolf pairs in relation to breeding season. Behavioral Biology 23: 521-525. Mech, L. D., L. G. Adams, T. J. Meier, J. W. Burch, and B. W. Dale. 1998. The wolves of Denali. University of Minnesota Press, Minneapolis. Murie, A. 1944. The wolves of Mount McKinley. U.S. National Park Service Fauna Series Number 5. Pulliainen, E. 1965. Studies of the wolf (Canis lupus L.) in Finland. Annales Zoologici Fennici 2: 215-259. Rabb, G. B., J. H. Woolpy, and B. E. Ginsburg. 1967. Social relationships in a group of captive wolves. Ameri- can Zoologist 7: 305-312. Rausch, R. A. 1967. Some aspects of the population ecol- ogy of wolves, Alaska. American Zoologist 7: 253-265. Schonberner, D. 1965. Observations on the reproductive biology of the wolf. Zeitschrift fur Sauzetierkunde 30: 171-178. Smith, D. W., K. M. Murphy, and D. S. Guernsey. 1998. Yellowstone Wolf Project. Annual Report YCR-NR-99-1. Yellowstone Center for Resources, Yellowstone National Park, Wyoming. Soper, J. D. 1942. Mammals of Wood Buffalo Park, northern Alberta and District of Mackenzie. Journal of Mammalogy 23: 119-145. Received 2 October 2000 Accepted 5 February 2002 Canada and the “Buffalo”, Bison bison: A Tale of Two Herds W. A. FULLER Professor Emeritus, University of Alberta; Adjunct Professor Athabasca University, Box 672, Athabasca, Alberta T9S 2A6 Fuller, W. A. 2002. Canada and the “buffalo”, Bison bison: A tale of two herds. Canadian Field-Naturalist 116(1): 141-159. From 1907 to 1912 the Canadian government purchased and imported more than 700 plains Bison, Bison bison, from Michel Pablo in Montana. A new national park, with an area of 159 square miles was established near Wainwright, Alberta, to accommodate them. It has generally been acknowledged that Buffalo National Park played an important role in saving the Plains Bison from extinction. This paper makes use of a packet of government files that were saved from destruction during the early1940s. The files deal mainly with events from 1912 to 1925, including the first appearance of bovine tuberculosis, and later the prevalence of tuberculosis in the herd. They also contain notes from the meetings of senior civil servants that led to the decision to transfer diseased plains bison to Wood Buffalo National Park, as well as summaries of submissions of those opposed to the transfer. One option, to slaughter the entire herd and start over with dis- ease-free stock, was rejected by well-meaning members of the public. When the Buffalo National Park was turned over to the military in 1940, 17 000 bison had been slaughtered as a result of annual culling. Ironically, had a total slaughter been carried out in 1923, fewer than 7 000 would have been killed. In addition, it is probable that we would have pure Wood Bison and no tuberculosis in Wood Buffalo National Park. In 1963, 18 disease-free Bison derived from a group of animals that showed some of the characteristics of Wood Bison, were released in the Mackenzie Bison Sanctuary. That herd now numbers about 2 600 individuals. As in 1923 we again have two herds, one with a high prevalence of tuberculosis and a second that is disease-free. In 1990 an Environment Panel (1990) recommended total depopulation of Wood Buffalo National Park and restocking with disease-free animals. As in 1923 the recommendation to slaughter and restock met oppo- sition on several fronts and so far no action has been taken. Must we repeat the serious error made in 1923? Key Words: Wainwright, Bison, tuberculosis, brucellosis, Pablo, slaughters, cattalo, transfer. More than thirty years ago I received from Dr. C. H. Douglas Clarke (see tribute and bibliography by Lumsden 1984a, 1984b) a box of old federal gov- ernment files that were related to the origin and sub- sequent operation of the Buffalo National Park near Wainwright, Alberta, and the transfer of Bison from there to Wood Buffalo National Park. According to a letter that accompanied the files, they were “...sal- vaged from the wartime file disposal in Ottawa.” Clarke gave as his reason for saving the files from destruction that “I realized that it was a personal record, kept by the late Maxwell Graham, to show that for his part he did his best to prevent the transfer of buffalo from Wainwright to the Wood Buffalo Park, and to promote herd reduction by slaughter.” Maxwell Graham was Director of Animals in the Dominion Parks Branch of the Department of the Interior from January 1912 until his transfer to a dif- ferent Branch on 1 January 1922. The files shed light on his role as guardian of the Bison in the Buffalo National Park near Wainwright, Alberta. At one time, those buffalo were much in the news, and some of the material in the files would have been “hot stuff’ politically. Clarke wrote: “Poor old Graham was in an uncomfortable spot, and I have a feeling of personal obligation towards him.” By accepting the files, I assumed that obligation, which I am only now attempting to fulfill. This paper will reveal information that has not been made public until now about the management of Buffalo National Park and about the transfer of buffaloes to the newly created Wood Buffalo National Park. My belief is that lessons learned so long ago are pertinent to the current situation in Wood Buffalo National Park. My hope is that those lessons will be taken to heart by all who have an interest in the long-term welfare of the buffalo.The files are the source of all direct citations set in small type and inset paragraphs. Plains Buffalo 1. Disappearance from the wild When Europeans first reached the great central plain of North America they were greeted by immense herds of “buffalo”, whose Latin name is Bison bison. Bison is now the approved common name for the beast, and I will use it from this point on. As time wore on and the human population increased, the Bison population decreased. The story of their decline has been told by several observers, such as Allen (1875), Hornaday (1889) and Isenberg (2000) for the United States; Hewitt (1921), Roe (1951) and Foster (1992) for Canada. Around the middle of the 19th century so many Bison had been killed that those remaining were essentially confined to two herds that were referred to as “southern” and “northern.” The southern herd was wiped out during the 1870s, and the northern herd followed in the 1880s. Fortunately, a number of ranchers, in both the United States and Canada, had captured and raised a 14] 142 THE CANADIAN FIELD-NATURALIST few Bison. One such herd is referred to either as the Allard, or the Pablo, herd. That herd was in Montana, conveniently close to Canada. 2. Recovery in Buffalo National Park After a considerable amount of negotiation, the Canadian government reached an agreement in 1907 with the owners of the Pablo herd to purchase a few hundred bison at $200 per head. They were to be delivered to a new park close to Wainwright, Alberta. However, the first shipment arrived before the fencing and buildings were finished in the new park, and the animals had to be released in Elk Island National Park, which had been set aside for the preservation of “elk” (American Elk, Cervus elaphus) in 1904. When Pablo began rounding up his animals, he discovered that he had more than the few hundred bison that Canada had originally contracted for. On 11 June 1907, 199 head arrived at Lamont,. Alberta, the rail station nearest to Elk Island National Park. On 11 October, another shipment of 211 head arrived at Lamont. No shipments were made during 1908 owing to illness on the part of Pablo, and the escape of 120 animals ready for transfer, but, during that year 73 miles (116 km) of fencing was put in place, and construction of other facilities was com- pleted at the new Buffalo National Park. In May of 1909, 325 of the Bison originally sent to Elk Island were recaptured and moved to the new park. The remainder of the 410 animals originally delivered in 1907, estimated to be only 48 head after the transfer in 1909 to Wainwright, formed the basis of the herd that still occupies part of Elk Island National Park. Later that year, Pablo delivered 218 head to Wainwright. In 1910, he was able to deliver 74 ani- mals and he made two shipments of seven bison each in 1912. In total, he was paid for 716 bison, 631 of which ended up in the Buffalo National Park. An additional 77 animals came from Banff National Park in 1909, and 30 came from the Conrad herd, which was also in Montana. According to one of the old files, the total number of animals introduced to the park was 738, but there is no record of the number of deaths or births during the years 1909 to 1911. Initially, the park, in total, occupied 159 square miles (a little more than 400 square km). Later, more land was acquired on which to grow hay for winter feed. Experts in the Depart- ment of Agriculture estimated that its carrying capacity was about 7000 bison. Vol 16 3. Events at Buffalo National Park Rate of Growth in Numbers I have been unable to find a complete set of annu- al census Figures, either in the old files or other sources, but I found nine numbers for the years 1911 to 1923 inclusive. (Table 1). Counts were apparently made in some, but not all years. Populations not under restraint tend to increase exponentially. A few hundred animals on a new range planned to support 7000 should be free of restraint. One of the properties of an exponential series is that the natural logarithms of the numbers lie on a straight line. The equation for the log, trans- formed data is LN(Nt) = 4.562 + 0.192 t where LN(Nt) is the natural logarithm of the number of Bison “t” years after their introduction to the Park; 4.562 is the logarithm of the number of Bison intro- duced to the park at time t = 0; and 0.192 (19.2%) is the annual rate of increase in numbers. The r? for this equation is 0.995. Entrapment of other species When the initial perimeter fencing was completed, park personnel discovered that they had enclosed wild elk, Mule Deer, Odocoileus hemionus, Moose, Alces alces, and a few Pronghorn Antelope, Antilo- capra americana (Lothian 1981: 29). I was suspi- cious about the antelope, but their presence was veri- fied by Rowan (1929: 359). Apart from competing with Bison for the available forage, there was a slight chance that the ungulates could have acquired diseases from contact with domestic cattle before they were trapped. In 1939, 1806 elk, 113 moose, and 242 deer that had been sharing the range with the Bison were slaughtered. Six per cent of the Moose and two individual deer “showed evidence of tuberculosis” (Lothian, 1981:37). Cross Breeding Experiments For several years The Department of Agriculture had followed the experiments of a Mr. Mossom Boyd in Ontario. Boyd had crossed Bison bulls with domestic cows and got fertile, hybrid offspring. The first generation hybrid offspring, when mated, also gave fertile offspring, which were named “Catalo” from the first three letters of “cattle” and last three letters of “buffalo.” The name was later changed to “Cattalo.” When Mr. Boyd died, the Department of Agriculture obtained his experimental animals and shipped them to an Agricultural Station at Scott, Saskatchewan. The agricultural scientists, anxious to do further experiments, entered into negotiations TABLE |. Available Counts of Bison in Buffalo National Park, 1911 to 1923. Year 191] 1912 1913 1914 Count 738 994 1188 1453 1915 1916 1919 1921 1923 1640 2077 3830 5052 6780 2002 FULLER: CANADA AND THE BUFFALO 143 with the Dominion Parks Branch to obtain some Bison and space in Buffalo National Park so they could conduct more experiments in cross-breeding. At this point, Maxwell Graham, Director of Animals in the Dominion Parks Branch, enters the story. The salvaged files contain a memorandum from Graham to Mr. J. B. Harkin, Commissioner of National Parks, dated 27th June 1916. Graham reminded Harkin that they had discussed the ques- tion of cross-breeding “prior to my departure for the west ... but the whole question was left in abeyance till Department of Agriculture was ready to take def- inite action.” He went on to write: I was only made aware the other day that such action had been verbally undertaken during my last absence from Ottawa, and subsequent action in confirmation thereof through correspondence has only now been brought to my attention. I am only now aware that it is proposed that a portion of Buffalo Park, about six square miles, be fenced off for the purpose of keeping therein the Cattalo herd bought by the Department of Agriculture from the estate of the late Mr. Mossom Boyd; That the shipment of these animals took place on or about November 26th last from Bobcaygeon Ontario, to Scott Saskatchewan; That certain other bovines, being cows of the Galloway breed, are also to be imported, I believe from the east, and to be added to the Cattalo herd for experi- mental purposes. I now desire to point out that if such action is taken, the herd of bison, now over 2000 in number at Buffalo Park, will incur considerable additional risk of becoming infected with some variety of infectious disease. Graham admitted that there was already “a certain risk to be faced of some infectious disease obtaining a lodgement in our main herd” but he pointed out that most of the land around the park was under cul- tivation and such land carries on it “but very few cat- tle of any kind.” He then expressed his fear that: We shall certainly run considerable risk, however, if we actually introduce into the park cattle in any quantity from Ontario. Some of these animals may, though them- selves immune, be carriers of disease from other herds. I would also point out that when an infectious disease is once brought into a large herd, the losses become very high, because it is difficult, if not impossible to check it after it has once obtained a foothold. He then went on to mention several such diseases — Hemorrhagic Septicemia. Anthrax, Foot and Mouth Disease — all of which had recently been discovered in the west. Finally, he most strongly recommended, in the same memorandum, that “some other area be found for the cross-breeding experiments... an area with natural drainage, and good running water and springs” instead of the “low land, stagnant water and liability to recurrent flooding of considerable areas” in Buffalo National Park. The Director of Agricultural Experimental Farms, J. H. Grisdale, lost no time in replying. His response is dated 5th July 1916 and required a little over three sheets of legal paper. His comments are in blue type, and the whole is peppered by comments in red type initialed M. G. (Maxwell Graham). The most impor- tant point comes near the end of Grisdale’s letter. I may conclude my remarks with respect to Mr. Graham’s memorandum by saying that after having perused it most carefully, and after having discussed it with the Veterinary Director General, with the Pathologist of the Department of Agriculture, each of whom assures me that, with average precaution, the probabilities of any infectious diseases being transmitted to your buffalo through the proximity of the herd of cat- talo are practically negligible. Graham’s response to that paragraph was directed to his own superiors. This, I consider completely absolves us from any pos- sible future criticism. Dr. Gordon Hewitt and myself each independently considered an added risk to be incurred & that is all I have contended. (signed) M.G. So the Bison, cattle, and cattalo of the Agriculture Department were established on 6 square miles of park land. Only a single fence, which permitted nose to nose contact, separated them from the Bison herd. Later, they were joined by several Yaks (Bos mutus). Grisdale guaranteed that all animals would be tested for diseases before being released inside the cattalo pen. Did any infectious diseases come with the experimental animals? We will never know for sure, but Graham was to raise the question again several years later. The cattalo episode did raise other questions. In a six-page (legal size) memo dated 7th July 1916 Graham reviewed the precautions already taken since 1912 and what was still needed. He made it plain that a major requirement was some means of segregating individuals by means of a system of cor- rals and cross fences. He had first made that request in 1914, but his request was turned down in a letter from Head Office to the Superintendent of Buffalo Park, dated 19th December 1914. The letter stated simply that “[T]he expense of constructing further fenced subdivisions would appear to be out of the question at present.” He fared no better in 1916. First Indications of disease The threat of disease was always a major concern for Graham because of his position as Chief of the Animal Division. His immediate superior, to whom he reported, was J. B. Harkin, Commissioner of Dominion Parks. There are a number of lengthy memoranda among the files from Graham to Harkin, urging that action be taken to reduce the risk of a serious disease outbreak. Virtually all of his plead- ing, such as the need for a system of corrals cited above, fell on deaf ears. It was almost 1917 before the first case of tuber- culosis was discovered. On 20 December 1916 a post mortem inspection of a dead bull revealed 144 lesions typical of tuberculosis. In January 1917 a visual inspection of the herd was ordered. Help was received from an inspector from The Agriculture Department. Out of the herd of 2 000+, only two or three animals were judged to be in poor condition. They were segregated from the main herd, but appar- ently not tested or slaughtered and examined post mortem. Veterinary Inspector Gillam wrote to Graham and recommended an enclosure with corral and a squeeze chute, “which would prove useful should it at any time be deemed advisable to vaccinate against Blackleg, Anthrax, or Contagious Abortion.” (The last of these is also known as Brucellosis.) No corral, let alone a squeeze chute, was forthcoming. No confirmed new case of tuberculosis showed up until 20 March 1919 when a dead cow was found, examined by the local Veterinarian, and diagnosed as having died from tuberculosis. The cow was thought to be about 15 years old. On 22 April 1919, a 4-year-old bull was killed and a post mortem examination revealed “traces of T.B.” On 13 April 1920, “an apparently healthy buffalo bull selected for slaughter for the Hudson’s Bay Company banquet, was found to be tubercular.” In spite of the evidence cited above, the Deputy Minister of the Department of Interior denied, in an interview published by the Montreal Gazette for 2 December 1922, that there was any tuberculosis in Buffalo National Park. Incontrovertible Evidence of Disease The first slaughter of Bison at Buffalo Park took place in February 1923, when 264 Bison were killed, butchered, and examined by a dominion veterinary inspector. Dr. A. E. Cameron (later the Veterinary Inspector General) and Dr. Seymour Hadwen, then of the Ontario Research Foundation, were also pre- sent. Cameron (1923) published a small paper enti- tled “Notes on Buffalo: Anatomy, Pathological Conditions, and Parasites” without even mentioning tuberculosis. Hadwen withheld publication of his observations until 1942. No details about disease are in the files that I have, but Lothian (1981: 32) wrote that “The inspec- tors found that 75 percent of the animals slaughtered (in 1923) had some form of tuberculosis lesion.” He did not mention, though, that the sample was strong- ly biased toward old males and thus was probably not representative of the herd as a whole. Lothian goes on to say (Page 32): “Dr. Hadwen recommended the elimination of the herd to avoid the spread of infection, but the proposal was not acceptable to park authorities.” The following win- — ter, 1923-24, 1847 bison were killed, but there is no record in the files of the occurrence of any disease. Dr. A. E. Cameron (1924) published a second paper entitled “Some Further Notes on Buffalo” In which THE CANADIAN FIELD-NATURALIST Vol. 116 he disposed of tuberculosis in a single sentence on the last page: “Tuberculosis has been found in buffa- lo, as is common in wild animals in captivity.” Lothian’s only comment was (page 32): “The presence of disease was withheld from public knowl- edge as a matter of departmental policy.” Apparently it was still a matter of departmental policy in 1981 when Lothian’s book was published. Instead of informing the public about the health of the herd, the government distributed pamphlets con- taining recipes for preparation and cooking of buffa- lo meat, whether from healthy or diseased animals. When Graham, who had transferred from the Parks Branch to the Northwest Territories and Yukon Branch at the beginning of 1922, learned the results of the slaughter he wrote a memorandum dated 13 April 1923 to W.W. Corry, Deputy Minister, Department of the Interior. It was marked “Personal and Con- fidential.” The pertinent points raised by Graham, with my comments in square brackets, were the following: As the official particularly charged with the adminis- tration of the buffalo in National Parks from January 1912 to January Ist, 1922, I am naturally concerned to hear that as a result of investigations made by Dr. Seymour Hadwen, he has reported the herd at Buffalo Park as being seriously infected with Tuberculosis I am fully aware that under the circumstances, once Dr. Hadwen’s report is made public, I, as former Director of Park Animals, must expect considerable crit- icism. Knowing as I do, and as our records will plainly show, that every effort to prevent such a calamity has consistently and all along been made by me, I personally do not fear such criticism. But the loss of your confidence in me I should feel very keenly, and in order to forestall such I wish to bring to your attention the following facts. Ist. During my tenure of office, so far as I am aware, there have only been four cases of tuberculosis, definite- ly so pronounced, in Buffalo Park. [The four cases already noted above. |] 2nd. During the slaughter of surplus buffalo in September 1922, Doctor Hadwen I understand, reports lesions in organs examined from the condition of which he estimated that not more than six years could have elapsed since infection took place. And, so I understand, infection in most cases was of a more recent date. 3rd. Our records will show that as far back as 1914, I strongly recommended that the Buffalo in Buffalo Park should be allowed to freely range during winter, instead of being confined in “winter quarters", and I stated “The danger of infectious or contagious diseases attacking the herd would not be as great (if allowed to winter in the main park) as if the herd were all kept together in con- fined quarters” (or confined feeding area.) My recommendation was turned down, without being referred to you, and it was not till July 1917 on the advice of Doctor Hargrave, Chief Inspector for Alberta, that action was taken to carry out my recommendation of 1914. I would suggest that Doctor Hadwen, as an unbiased authority, might be asked his opinion on the points 2002 raised in my memorandum of June 27th, 1916, and whether or not in his opinion the recommendations made in my memorandum re tuberculosis of March 19th, 1919 were sound. In any case, these latter were not acted on except in so far as reference is made to them in the attached memo- randum of April Ist, 1919 by Doctor Torrance. The part of Dr. Torrance’s letter to which he refers seems to be the following: The whole question is one of providing plenty of nourishment and fresh air. The buffalo, I understand, are constantly out in the open, and they should always be provided with an abundance of nutritive food so that they may be maintained in as vigorous a condition as possible. The next, and last, point raised by Graham must be read in association with his 2nd point. It only remains to be added that my attempt in 1916 to prevent what I then considered, and still consider, an uncalled for additional risk to the buffalo...was met with by unmerited censure and ridicule. When in September 1922 Doctor Hadwen made his investigation of the herd in Buffalo Park, just six years had elapsed since, against my protest, the cattalo and domestic bovines were first placed in Buffalo Park. Graham is clearly making a connection between arrival of the Bison, cattle, and yaks in 1916 and Dr. Hadwen’s finding that none of the cases of tubercu- losis were older than 6 years, i.e. acquired no earlier than 1916. He does not go quite so far as to make an outright accusation. Without, however, actually blaming the cattalo and domestic bovines in the experimental herd for the seri- ous condition of our buffalo in Buffalo Park, it would seem that my modest expression of opinion as to the added risk incurred by insistence in selecting the place for cattalo experiments where it is was fully justified. Written in pencil across the upper left hand corner of the first page of the memorandum is the following: “Mr. Graham, I don’t see how you can be blamed in this matter. W.W.C.” The initials are those of W.W.:Cory, Esq., C.M.G., Deputy Minister, Department of the Interior, to whom the memoran- dum was addressed. There is also a short memorandum directed to Mr. Graham and signed by O. S. Finnie, Commissioner of the Northwest Territories and Yukon Branch, Department of the Interior, to which Graham now belonged. It read: I read over, with much interest, the correspondence hereunder. The evidence shows conclusively that you always entertained apprehensions as to the safety of the herd at Wainwright — even in the face of expressions to the contrary from those in high authority. They cannot lay at your door the charge that you failed to advise them of the dangers of disease. I interpret this to be, on the part of the government at least, a clear exoneration of Maxwell Graham. The Horns of a Dilemma Even before1920, the relentless pressure of expo- nential increase in the size of the herd, coupled with FULLER: CANADA AND THE BUFFALO 145 deterioration of the range, created a serious problem for Buffalo Park managers on the site, senior civil servants in Ottawa, and ultimately, elected politi- cians. Something had to be done — but what? The only options appeared to be to continue with annual slaughters, or slaughter of the entire herd, or transfer the surplus to some other suitable location. Hewitt (1921: 136), whose book was published posthumously, had some further suggestions. One was to “establish small parks in other parts of the Prairie Provinces, where small herds could be main- tained, which would make the bison accessible to people’. He suggested also (136) “[E]very large city should have its zoological park.” And finally he set out his view on the possibility of domestication. “The greatest value of the buffalo, however, lies in the possibility of its domestication... the greatest need of the prairie provinces is an increase in its beef-producing capacity. The buffalo is an animal which offers great possibilities, being pre-eminently suited to prairie conditions, and at the same time it produces a robe of no small commercial value” (136). At the time when this was written, no one seemed to take domestication seriously, and disease did not seem to be a barrier. Today, in North America, there are many more Bison on private ranches than in parks and reserves. When the slaughter of nearly 2000 bison in 1923 was made public there was a strong reaction against further slaughters. Given the strong public objection to a simple herd reduction, the Parks Branch felt that total elimination of the Buffalo National Park herd, as suggested by Hadwen, was out of the question. Dr. C. H. D. Clarke, who salvaged the files, was also in favour of total slaughter and restocking with healthy Bison from Elk Island National Park. Wood Bison In January 1772 Samuel Hearne and his Indian companions crossed Great Slave Lake from north to south. Hearne (1795: 250) wrote: “Immediately on our arrival on the south side of the Athapuscow [now Great Slave] Lake, the scene was agreeably altered, from an entire jumble of rocks and hills, for such is all the land on the North side, to a fine level country, in which there was not a hill to be seen, or a stone to be found.” As they made their way up the lowlands on the eastern side of the Slave River, he noted that “Buffalo, moose, and beaver were very plentiful....” Hearne (1795: 251) also noted that “The buffalo in those parts, I think, are in general much larger than the English black cattle: particularly the bulls, which, though they may not in reality be taller than the largest size of the English oxen, yet to me always appeared to be much larger. In fact, they are so heavy, that when six or eight Indians are in company at the skinning of a large bull, they never attempt to turn it over while entire, but when the upper side is 146 skinned, they cut off the leg and shoulder, rip up the belly, take out all the intestines, cut off the head, and make it as light as possible, before they turn it to skin the under side. ....The head of an old bull is of great size and weight indeed: some which I have seen were so large, that I could not without difficulty lift them from the ground.” Hearne was thus the first European to see and describe the wood buffalo. When I was at Fort Smith in the 1940s and 1950s I took part in the skinning of many a Bison, and I watched many Indians and Métis working in pairs. I never saw them lighten the load before turning over a Bison in Wood Buffalo National Park. Hearne was not usually given to exaggeration, as were many explorers of his day, so I take his obser- vations as evidence that the Wood Bison were con- siderably larger than those of the plains. Other early explorers saw Wood Buffalo in the course of their journeys. While visiting Forts Chipewyan and Wedderburn, on Lake Athabasca, Captain John Franklin (1823:154) noted that “The traders here also get supplied by the hunters with buffalo and moose deer meat (which animals are found at some distance from the forts)...” After descending the upper Slave River, and nego- tiating the rapids above the present Fort Smith, Franklin ascended the Salt River as far as the salt springs, where his men collected salt for the rest of their journey. On leaving the springs (Page 107) they “perceived a buffalo plunge into the river before us.” The unfortunate animal was immediately shot and kept the entire crew in meat until they reached Fort Resolution on Great Slave Lake. While there he was informed (199) that the residents “Procure moose, buffalo, and rein-deer meat occasionally from their hunters; but these animals are generally found at the distance of several days’ walk from the forts.” Joseph Sabine, a zoologist of Franklin’s time, pre- pared The Zoological Appendix (Number V), to Franklin’s narrative. About the Bos Americanus or American Buffalo, he wrote (1923: 688) “... they are extremely numerous on the plains of Sashatchawan,(sic) and are also found, though less plentifully, in the woods as far north as Great Slave Lake; a few frequent Slave Point, on the north side of the lake, but this is the most northern situation in which they were observed by Captain Franklin’s party.” The reference to Slave Point, on the north side of the lake, has been cited by several other authors, who, apparently, did not verify its location. Slave Point was said to be composed of limestone. The only Siave Point on modern maps is at approximate- ly 61° 10' N and 115° 55' W on what I would call the west shore rather than the north shore. No Slave Point appears on Franklin’s map on the true north shore which, in any case is composed of Precambrian granites, not flat limestone. THE CANADIAN FIELD-NATURALIST . Voi. 116 In the middle of February 1890, Warburton Pike (1892: 143) set out from Fort Resolution on a buffa- lo hunt with an Indian guide named Francois. His assessment of the distribution of bison was as fol- lows: “Scattered over this huge extent of country are still a few bands of buffalo. Sometimes they are heard of at Forts Smith and Vermillion, sometimes at Fort St. John close up to the big mountains on Peace River, and occasionally at Fort Nelson on the north branch of the Liard.” After reaching the end of Francois’ road, and walking until noon, they came on a band of eight bison and killed one cow. A few weeks later he made a second hunt, again with Francois as his guide, but they saw no trace of bison. Pike (1892: 43) appears to have been the first to attribute the size of the wood buffalo to environmental factors as the following quotation shows: “These animals go by the name of wood buffalo and most people are of opinion that they are a dis- tinct race from the old prairie buffalo so numerous in bygone days; but I am inclined to think that the very slight difference in appearance is easily accounted for by climatic influences, variety of food, and the better shelter of the woods.” The italicized words are one of the untested hypotheses that will play an important role in what happened later. In 1892, Frank Russell, a young academic at the University of Iowa, began his journey into the far north. On 9 January 1893, Russell set out from Fort Resolution, guided by Francois who had guided Pike. As Pike had done, he began his journey on the Little Buffalo River and later turned off to the south- west. On the fourth day they reached what he called (Russell, 1898: 103). “the northern limit of the buf- falo range, perhaps fifty miles south of the Great Slave Lake.” They saw no buffaloes and had to return empty handed after 13 days on the trail. Early in the spring of 1902, Charles Camsell was asked to search the country south and west of Fort Smith for Wood Bison. At that time he was a very junior member of the Geological Survey, which was in the Department of the Interior. Before he retired, at age 70, he had risen to Deputy Minister of the Department of Mines. His journey is set out in his memoirs, Camsell (1954). He and his assistant launched their canoe at Athabaska Landing (now Athabasca) and paddled down to Smith Landing (now Fort Fitzgerald). From there they went by ox cart to Fort Smith. From Fort Smith, they made two short sorties and saw only one small bison herd and many tracks. Their final trip took them to the Peace ‘River by way of the Salt River, the upper Little Buffalo, and the Jackfish River. They began that journey on 2 August and reached the Peace a whole month later. Camsell estimated that the number of Wood Bison was probably about 300, but he gave no data of any kind to support his estimate. 2002 FULLER: CANADA AND THE BUFFALO 147 Ernest Thompson Seton and Edward A. Preble set out for the north, by canoe, in 1907. Seton could not have chosen a better companion than Preble who, with his brother, covered about 1200 miles by canoe in the Hudson Bay region in 1900, and in 1903-04 travelled several hundred more miles in the Athabaska-Mackenzie Region. En route to the tundra, Seton and Preble paused at Fort Smith for a buffalo hunt. Their guide led them to a group of 13 bison the first day (page 48) and 22 the second day (page 52). An account of Seton’s trip was published in 1911 (Seton 1911) and republished in 1943. It is apparent from this brief survey that Bison lived at least as far north as the south shore of Great Slave Lake and perhaps further. It is also apparent that by the end of the 19th century, their numbers were rather small, and the largest herd was thought to live in the angle formed by the junction of the Peace and Slave Rivers. Maxwell Graham visited the area south of Fort Smith in 1912. On his return to Ottawa he wrote a memorandum, dated 16th October 1912 to J. B. Harkin. He dealt briefly with the character of the country through which bison ranged, the length of time that they are known to have lived in the region, and the size and type of “these pure-blood bison.” He argued that although ample range is critical for their survival, ample range alone is not enough. They also need other kinds of protection. Graham suggested that protection would be easier and less expensive if the various small groups of bison could be herded into a single reserve. He felt that the ideal location for such a reserve was the junction of the Peace and Slave Rivers (Figure 1). He noted that: A line drawn eastwards from Peace Pt. upon the Peace River, would intersect the Great Slave River about 20 miles below the junction with the Peace. If a suitable fence were constructed along this line, a distance of 25-30 miles, the Buffaloes could gradually be worked from the North into this peninsula where there would be ample range for a number of years. Graham thought that “herders” could turn back any Bison that sought to cross the Slave or the Peace River in winter over the ice, but he neglected the possibility of escape by swimming across the rivers in summer, which Bison do with ease. Graham also quoted from what he called the “Smithsonian report.” (See Hornaday 1889). “There is reason to fear that unless the government takes the matter in hand and makes a special effort to prevent it, the pure-blood bison will be lost irretrievably through mixture with domestic breeds, and through in- breeding; this latter causes eventually, loss in size and final sterility.... As I have already stated no suitable stock can now be procured from any of the herds in the United States, a large number of these are not pure- blood, some which are, have deteriorated in size owing to too close confinement and in-breeding,” That most of the herds in the United States are not pure-blood has recently been confirmed by mito- chondrial DNA sequencing (Polziehn, et al., 1995.) Graham’s memorandum closes with the following paragraph. Such being the case I again urge that the future administration of this Northern Canadian Herd, the last and best of the buffalo, be entrusted to this Dominion Parks Branch, and that provision be made in the esti- mates to provide for the enclosure proposed above, a plan of which is appended herewith and that a vigorous policy as to this herd’s protection be instituted, and a capable superintendent be appointed who shall have the interests of this Branch at heart, and who would also understand and appreciate the policy of the Dominion Parks Branch.” No action was taken on Graham’s memorandum. The need for a reserve of some kind was raised again in 1914 at a meeting of the Council of the Northwest Territories, which met in Ottawa in those days, but again it was not acted upon. Camsell made three more short trips into the Wood Bison range during the summer of 1916 at the request of the Parks Branch of the Interior Depart- ment. A type-written copy of his report is among the salvaged files. On page 1 Camsell wrote: The wood bison of northern Alberta and the adjacent portion of the Northwest Territories are in two separate bands occupying two distinct ranges, and there does not seem to be at present or within recent years any migra- tion of buffalo from one range to the other.... A belt of muskeg country 30 to 40 miles wide separates the north- ern from the southern range and prevents migration from one herd to the other except by way of the Salt plain. The italicized portion is the second untested hypothesis that was to play a role later on. The second important conclusion in Camsell’s report is that most of the Wood Bison were in an area extending from the Peace River north to latitude 65° (clearly a typographical error) between longi- tudes 112° and 113°. On page 5 he suggests a game reserve bounded by 60° on the north, 113° 20' on the west, latitude 59° 10' on the south, and Slave River on the east. No action was taken on his recommen- dation for a reserve. In 1918, another recommenda- tion to create a park was also turned down. In 1922, Maxwell Graham revisited the range of the Wood Bison. His report, entitled “Some obser- vations on the wild buffalo in Northern Alberta and the Fort Smith District, Northwest Territories,” fills 5 sheets of legal paper and is dated 23 November 1922. The report begins with a notation that “true buffalo” are found only in Africa and Asia, and that the “buffaloes” of Europe and North America are properly known as “bison.” There is only one species of Bison in Europe, Bison bonasus. There is also only one species in North America, Bison bison, which was thought by many to have two subspecies. a — 148 THE CANADIAN FIELD-NATURALIST Graham wrote: Both the writer and his colleague are in a position to state that our Northern wood-buffalo are larger, darker, and handsomer specimens than those which formerly ranged the plains far to the south. He goes on to say, however, that: in the writers’(sic) opinion, whatever differences there are between it and those of the plains, is (sic) entirely owing to environment. Graham clearly agreed with Pike’s untested hypothesis. It is not clear whether or not Graham believed that Wood Bison would revert to Plains Bison if they were moved to central Alberta, or con- versely, Plains Bison would come to resemble Wood Bison once they were moved to the wood bison envi- ronment. On page 2 of the memorandum Graham wrote: The buffalo at Elk Island Park are supposed, in the aggregate, to be the best in any park, nevertheless, with- out meaning to disparage this fine herd, the writer and others who were with him on the wood-buffalo range must state that in size, thriftiness, stamina and general appearance the wild wood-buffalo is in a class by itself and quite unapproachable. The use of “in a class by itself’ suggests that they were worth preserving whether or not they were for- mally recognised as a subspecies. Graham made his headquarters at Pine Lake in 1922. A wagon road connected Ft. Smith and Pine Lake. About two miles before it reached the lake, the trail crossed the upper Salt River. When I worked in WBNP in the late 1940s, the old trail and the cross- ing were still visible and I visited the old crossing, and saw the tree in which Graham had a platform from which to watch Bison. One of the first “buffalo rangers,” Billy McNeil, was my informant. Here are some excerpts from notes Graham made on August 12, 1922. The most successful day yet. Dull and raining in the morning, but clearing towards noon. Stewart and I start- ed (on foot) for Salt River Crossing. At Wallows on the way I spotted a huge bull lying down, however, he wind- ed us and got away before we could get a picture. On arrival at Salt River Crossing, Stewart climbed a leaning tree and I posted myself across the river under a small tamarac,(sic) in full view of open muskeg. We saw and got pictures first of a herd of 16 buffalo. In about half an hour another herd of about the same size came out of the forest, of which we also got some pictures. Stewart fol- lowed these in hopes of getting some close-ups, none of them having seen or scented us...Shortly after, Stewart returned not being able to catch up the buffalo in the bush beyond river (sic), and it was then that I heard more coming... So far as I could estimate there were 9 in this herd, one immense bull, two cows, two 2 year olds, 3 yearlings and | very small calf, whose skin was throughout of a faint yellowish white colour. All in all, Graham and his assistant saw and counted 114 bison, which was more than all other visitors mentioned above had seen. He also ventured to estimate the total number. His estimate was based upon Vol. 116 ...the finding of old trails and old wallows being reopened as well as the presence of new ones, and from the number of young stock seen it is safe to state that the wood-buffalo are increasing, and from all the evidence our party was able to investigate, at a conservative esti- mate, the number of buffalo in the southern range may be put down as 1000. Graham also argued again for more protection from poachers, forest fires, and Wolves (Canis lupus), in the form of an enlarged corps of rangers (now wardens). The last paragraph in the report sug- gests that Graham had some inside information. Both for the protection of the buffalo as well as valu- able forms of other wild life, it is expected that steps will shortly be taken to conserve for posterity the range on which the buffalo roams. Another memorandum dated 13th November 1922 is entitled “Reasons Why the Wood Buffalo of Northern Alberta and Northwest Territories should be preserved.” These buffalo are the last of their species living to- day under absolutely free and wild conditions. They are the finest specimens of their species, superi- or in pelage, size, and vigour to those of the plains, whose descendants to-day exist in our parks. Owing to certain physiological causes buffalo in cap- tivity deteriorate in size and stamina, further, when in confinement their natural increase contains an undue proportion of males to females, which unless steps are taken to remedy this surplusage (sic) of males is apt to bring about tuberculosis and kindred diseases. We know now that sex is determined by presence or absence of a Y chromosome, and it is not clear how the sex ratio is involved in the spread of dis- ease. The time is approaching when an infusion of new, unrelated, blood will be needed by our herds in the National parks, and it is only from the northern herd that such infusion can be obtained. Looking to the future success of the experimental cross-breeding between buffalo and domestic bovines, it is imperative that a reserve stock of pure blood bison of the highest potency should be kept in reserve, so that the ultimate fixed type of new range animal may continue to pass on to successive generations the prepotent qualities of the true bison, viz hardiness, thriftiness, a valuable robe and first class beef qualities. A second typescript is also among the files. It may be a draft of “Canada’s Wild Buffalo,” which was published by the Department of Interior in 1923. It contains more information than the memorandum about the nature of the country, such as meadows, muskegs, sloughs, poplar ridges with edible grasses, jack pine ridges that have a scanty ground flora, and sink-holes that develop in the gypsum that underlies much of the area. Associated with the report and manuscript was a letter, dated 22 November 1922, to Graham from R. M. Anderson, who was then Chief, Division of Biology, Victoria Memorial Museum. His letter reads, in part, as follows. 2002 FULLER: CANADA AND THE BUFFALO 149 FIGURE 1. Maxwell Graham’s original map showing the area he proposed as a reserve for wood buffalo in the angle between the Peace River, which comes in from the southwest, and the Slave River which exists to the north. The line near the top is the 60th parallel. The grey patches in the lower right are water bodies. I have carefully read over the manuscript of your description of the wood buffalo range, observations on the habits of the wood buffalo, and reasons why the wood buffalo of northern Alberta and Northwest Territories should be protected. It is very interesting as a very up-to-date summary of the wood buffalo situation as well as a comparison of the wood buffalo with the more southern herds, and there is really nothing that I can criticize in the paper. The desirability of keeping a reserve stock of pure 2 ee 6 ee ees a 150 blood buffalo, living under free, natural conditions, is so obvious as to need no argument.... Thanking you for your kindness in loaning me the report, I remain Yours sincerely, (signed) R. M. ANDERSON About one month after Graham submitted his reports, the steps that he had predicted came to pass. An Order in Council established Wood Buffalo National Park (WBNP) in December 1922. The stage was now set for the two streams we have been follow- ing to come together. Suddenly, 10 500 square miles of sparsely populated wood bison range became avail- able to receive the excess in the National Park at Wainwright. The Transfer Goes Ahead The following information is taken from a docu- ment assembled by Col. J. P. Richards dated 6th May 1923. Col. Richards held a senior position in the Department when I joined it in 1947, and I found him to be an honest, dedicated public servant. The title of the document is “Summary of correspon- dence dealing with the transfer of the buffalo from Wainwright to Wood Buffalo Park.” Notations in square brackets are my comments. 1. The decision makers Under date of 23rd May, 1923, the Commissioner of Canadian National Parks advised the Department that there were too many buffalo in Wainwright Park and some arrangement would have to be made to reduce the number. The Department suggested that instead of slaughter- ing excess buffalo it might be desirable to transfer some of the healthy [?] young stock to the Wood Buffalo Park and it was arranged to call a conference to discuss the proposal. The conference was held on the 30th May and the following extracts which are taken from a memoran- dum dated 5th June, 1923, outline the decisions arrived at. Under date of May 30th. 1923, on Wednesday at 3 p.m. a conference was held at the Commissioner’s office, at which were present Mr. W. W. Cory, acting in a dual capacity as Deputy Minister of the Department and Commissioner of the Northwest Territories, Mr. O. S. Finnie, Director of the Northwest Territories, Mr. J. B. Harkin, Commisssioner of National Parks, Doctor F. Torrance, Veterinary Director General, Department of Agriculture, and Mr. A. Smith, Superintendent of the Wainwright Park. It is important to note that Maxwell Graham’s name is absent from the list of participants. He took no part in that conference. The question before the Conference was concerned with the disposal of buffalo surplus to the capacity of the Wainwright Park, and also the advisability of introduc- ing these surplus buffalo into the Wood Buffalo Reserve. The question was rendered more complex owing to the fact that tuberculosis had manifested itself among the buffalo in the Wainwright Park. [My emphasis] THE CANADIAN FIELD-NATURALIST Vol. 116 The conference was opened by the Deputy Minister’s statement that Mr. Harkin had informed him something must be done to relieve the growing congestion of buffa- lo at the Wainwright Park, that the slaughter of a large number of buffalo had been suggested, but he disliked the idea of such slaughter and had thought a way out of the difficulty could be found by transplanting the surplus Wainwright buffalo into a new environment away from all danger of communicable disease. [Except what they took with them, of course.] . Such an area was to be found in that area selected by the buffalo themselves, whose descendants are now known as wood buffalo and whose habitat near Fort Smith has recently been created a park. Addressing in particular Doctor F. Torrance, the Deputy Minister requested his opinion with respect to the chances of recovery from tuberculosis if buffalo infected with this disease were transplanted in the Fort Smith district, and if in his opinion there would be any serious menace to the wood buffalo through such action being taken. Doctor Torrance was of the opinion that although in some less advanced cases an improvement might be looked for, the transplantation of any considerable num- ber of the infected herd into the range of one, presum- ably, not infected would be extremely hazardous. It was then finally decided that selected young stock, not over one year old, should as soon as possible, be seg- regated, corralled, and squeeze-chuted, in order that each one might be subjected to the intradermal test. Those passing the test to be shipped by rail to Peace River Crossing, thence by scows, through the [Vermilion] chutes, to Peace Point and liberated in the Wood Buffalo Reserve. [Peace Point is an outcrop of limestone at least 15 metres high!] Dr. F. Torrance of the Department of Agriculture, who was present at the conference, expressed his opinion about the disease problem in a letter to the Department on the 30th of the same month. As regards Tuberculosis, the suggestion has been made that some of the younger buffalo should be trans- ported to the range of the wood buffalo in the north and permitted to mingle with them. This proposition is objectionable from a health point of view, in that it would be almost certain to carry infection to this herd of wood buffalo, which presumably is at present free from this disease. [My emphasis] If this proposition were, however, modified and prep- aration made so that young animals up to the age of yearlings only were transferred, and that these animals were, previous to transference, submitted to the tuber- culin test, so as to eliminate any that reacted, much of the objection would be removed. Note that Torrance did not give his outright bless- ing to use of the tuberculin test. As late as the 1950s at least, the tuberculin test gave some false negatives so Dr. Torrance would have known that a single test would not have detected all reactors. Under date of the 13th November, 1923, the Edmonton Journal published a despatch from Ottawa in which it was stated that the Minister of the Interior had announced that it was the intention to transport two thousand buffalo calves from Wainwright to Wood Buffalo Park. 2002 A second Parks Branch Memorandum dated 20th November, 1923, dealt only with the steps to be taken to prevent injury to the animals in transit from Wainwright to WBNP. The Commissioner, Canadian National Parks Branch, on the 22nd of the month, commented upon the memorandum referred to in part as follows: There is one subject which has been discussed in the department which is not mentioned in Mr. Nagle’s mem- orandum, that is the possibility of the transfer of Wainwright buffalo to the wood buffalo range resulting in the introduction of tuberculosis in the wild herd. You will remember that last spring we had a conference with Dr. Torrance upon this subject and that the Doctor would not commit himself concerning this aspect of the situation. A letter dated 27 November 1923 from J. B. Harkin to Edmund Seymour, President of the American Bison Society was published in the Report of the American Bison Society for 1922—23 page 32. It read as follows: “Since last winter the Department has had under con- sideration a proposal to make shipments of buffalo from the surplus of the Wainwright herd to Fort Smith coun- try, now occupied by the so-called wood bison. It is the intention of the department to make some experimental shipments next Spring but I doubt very much whether the number will be as large as one thousand. Difficulties in connection with the transportation of adult buffalo to such a remote area are so great that it is improbable that any attempt will be made to send adults to the north. Yearlings appear to be the only type that could be han- dled on such a project. Just what number will be sent has not been decided. I do not think a decision can be reached until next spring. The only established fact in connection with this subject is that the Department’s present intention is to make an experimental shipment.” Nowhere does the summary of discussions on movement of Bison mention an “experimental ship- ment.” So why did Harkin write the letter? It could have been to alert Seymour with the hope that he would protest the shipment in the name of the Bison Society. It is even possible that Maxwell Graham drafted the letter for Harkin’s signature. The copy of the Bison Report for 1922-23 that arrived with the files has Graham’s name and departmental address on the cover. There followed a copy of a report by an Inspector Waddy on the prevalence of tuberculosis in 46 bison. According to the file that lists all slaughters from 1923 to 1937 there was “no slaughter” in 1924-25, but a copy of the report, dated 10th Jan’y 1924, was sent to the Veterinary Director General, and a copy is among the salvaged files. The report begins with an arbitrary, but adequate, definition of three age classes. The definitions are as follows: With not more than one pair of permanent teeth up — under 5 years With all permanent teeth up but not worn at the edge — 5-10 years With all teeth in wear — over 10 years. FULLER: CANADA AND THE BUFFALO 151 TABLE 2. Prevalence of tuberculosis in a sample of 46 bison examined by Dr. Waddy in January, 1924. Age Number Number Percentage Class Slaughtered Positive Positive 1 15 2 13.3 2 13 9 69.2 3 18 16 88.9 TOTAL 46 4 | 58.7 Table 2 shows Dr. Waddy’s findings. Waddy remarked “I think this plainly shows the prevalency of generalized T.B. in the old animals.” The numbers certainly speak for themselves. Prevalence increases with the age of the animal, and more than half of the animals in the sample were infected with tuberculosis. The findings also help to explain the high incidence of tuberculosis in the 1923 slaughter that was biased toward old males. The Richards memorandum went on as follows: This report was received in the Department at the time consideration was being given to the proposal to transfer a number of Wainwright Buffalo to Wood Buffalo Park. Jt will be observed that fifteen animals five years of age and younger were examined by Inspector Waddy and only two showed slight evidence of tubercu- losis. No animals over the age of three years were trans- ferred to Wood Buffalo Park and the number of three- year olds that accompanied the shipment was less than five per cent of the total. At a conference held in the Deputy Minister’s office on the 9th April, 1924, it was decided to transport 500 yearling buffalo from Wainwright to Wood Buffalo Park during the summer of 1924. It was decided that the Parks Branch would assume responsibility for segregat- ing, corralling and squeeze chuting and placing the buf- falo in cars at Wainwright, that the Northwest Territories Branch would be responsible for the care and transfer of the buffalo from Wainwright to Waterways, thence by barge to Wood Buffalo Park. This arrangement was con- curred in by the Minister under date of 24th April, 1924. Instructions respecting the shipment of the buffalo were received by the Superintendent at Wainwright too late to permit of the young buffalo being segregated in time for shipment in 1924 and it was decided not to attempt to transfer any buffalo until the following year. The Superintendent of Wainwright Park was given instruction to make plans for the transfer of at least two thousand young buffalo in 1925. [Note that it is now “young” buffalo, not yearlings. | In [a] memorandum dated 6th October, 1924, the Commissioner of the Canadian National Parks refers to a conference which was held in the Deputy Minister's office on the 3rd idem when the following decisions with respect to the transfer of the buffalo from Wainwright to Wood Buffalo Park were made: 1. In 1925 at least two thousand buffalo would be shipped. 2. The shipment would include only one-and-two- year old animals from the 1923 and 1924 increase. 3. That the tubercular test would be dispensed with. [My emphasis] 152 4. That the responsibility of the Canadian National Parks Branch would cease when the animals were load- ed on the cars at Wainwright. If we take the Waddy report at face value, that is two tubercular animals out of every 15, then around 900 of the 6673 animals delivered would have been carrying tuberculosis. The actual ages and numbers transferred over the next four years are shown in Table 3. Another column in the file listed deaths en route, which totalled 47 animals accidentally crushed or injured. None of the reported deaths “were attributed to tuberculosis.” The next sentence, however, reads: “The carcasses were not examined to ascertain whether the animals were infected, since there was no government official’ available qualified to make such an examination.” My home in Fort Smith in 1947 was no more than 100 m from the home of an elderly gentleman who had been the Chief Warden of Wood Buffalo National Park when the transfers were made. He told me that many animals died as they walked down the gangplank at the landing, or as they made their way along a trail that ran for about 2 miles from the river through some woods to the nearest patch of prairie. Other old timers at Fort Smith, who had taken part in the unloading, confirmed the Chief Warden’s story. When I mentioned this conversation in a letter to Ottawa in 1947 I got a reply that said in no uncertain terms that 6673 animals were delivered, apart from 47 that died en route. Apparently, the translocation was still a touchy subject 20 years after it happened. The next paragraph in the Richards memo lists the names of several people who were opposed to the ven- ture. Another document lists arguments pro and con. 2. Controversy about the translocation Two things that caused the most concern among opponents of the transfer were first, that infectious disease might be introduced to the Wood Bison, and second, the Wood Bison would disappear as a sub- species through hybridization with the introduced plains bison. Those who defended the transfer based their argu- ments mainly on three untested hypotheses. First came Camsell’s claim that the northern and the southern herds of Wood Bison were separated by a band of marsh and muskeg that could not be crossed. THE CANADIAN FIELD-N ATURALIST Vol. 116 Thus, regardless of what happened to the southern herd, the northern herd would not hybridize and it would not be exposed to disease. Second came Pike’s belief, supported by Graham and others, that any differences between plains and Wood Bison were superficial and due to environmental factors. Some supporters of this view went so far as to pre- dict that the introduced animals would gradually develop the characteristics of Wood Bison. Third, the introduced animals were too young to carry any infectious disease including tuberculosis. Both J. B. Harkin and Dr. Torrance, however, expressed con- cern after the rejection of Dr. Torrance’s recommen- dation, that only yearlings that passed a tuberculin test were to be shipped. Among the defenders was Maxwell Graham, even though he was no longer associated with the Dominion Parks Branch. In a one page Letter to the Editor of The Canadian Field-Naturalist (Graham 1924) he upheld Camsell’s belief in the barrier between northern and southern herds, and the belief that the difference between plains and Wood Buffalo was just a matter of different environments. I believe that he genuinely believed that the transfer “will be the means of saving for posterity the calf crop at the Wainwright Park for 1922-23 and succeeding years.” Mr. Edmund Seymour, President of the American Bison Association, and Dr. W. T. Hornaday, an American mammalogist, met with the Deputy Min- ister. The stance taken by the Bison Association would have carried considerable weight among the general public. However, the Deputy Minister per- suaded Seymour and Hornaday to give their approval to the actions taken by the government. The warning they received in Harkin’s letter, if it really was a warning in disguise, was ignored. On the other hand, a considerable list of people, many of them well versed in mammalogy, expressed their objections directly to the government or through published articles. Dr. D. Rutherford, Department of Agriculture, University of Saskatchewan, told the Minister in a letter that the transfer was a mistake. W.E. Saunders (1925), a naturalist living in London Ontario, wrote “it would be better to lose the entire Wainwright herd, rather than risk the last rem- nant of the Wood Buffalo.” TABLE 3. Numbers and Ages of Bison Transported from Wainwright to Wood Buffalo Park in the Years 1925 to 1928. Season Yearlings Two Years Three Years Total 1925 1127 507 - 1634 1926 1435 493 83 2011 1927 1255 436 249 1940 1928 1009 79 - 1088 TOTAL 4826 1515 332 6673 eee 2002 FULLER: CANADA AND THE BUFFALO 153 Professor A. B. Howell, Chairman of the Ameri- can Society of Mammalogists, cited a resolution passed at their Annual Meeting condemning the transfer because “distinctive characteristics of the Wood Buffalo would be lost in a few generations,” and “TB and other diseases would be likely to be transmitted with harmful effects to the northern herd.” Dr. Francis Harper (1925: 45), an American mam- malogist who had travelled from Lake Athabasca to Great Slave Lake with Charles Camsell, and who had visited the Park in 1920, made a number of points in a letter to the Editor of The Canadian Field-Naturalist. First, “The wood buffalo (Bison bison athabascae) is too important an animal to be subjected to experimentation that may result in decided harm to the entire subspecies.” Second, “The proposal outlined by Graham raises the old question of man’s interference with nature, which, in too many cases is alike unnecessary and unjustifi- able.” Third, “Establishment of the Park was one of the most important and far-sighted conservation measures ever adopted by the Dominion Govern- ment.” The implication is why spoil it? Fourth, with respect to the separation of the northern and southern herds he wrote “This gap can hardly yet be accepted as a proven fact, or as a necessarily permanent con- dition. There is evidently no physical barrier that would prevent the two herds from mixing.” Fifth, “The possible transmission of disease through the introduced plains buffaloes is another factor to be considered.” Sixth, if the Wainwrights cannot be set- tled in Alberta, “would it not be wiser to send them to the slaughter-house at once?” Two members of the National Parks Branch faced censure over publication of Harper’s critical letter to the editor. One was Mr. Hoyes Lloyd, Supervisor of Wildlife Protection, who was also President of the Ottawa Field Naturalist Club at the time. The club’s journal was, and still is, The Canadian Field Natur- alist. The Editor of the journal was Dr. Harrison F. Lewis, who later became the first Director of the Canadian Wildlife Service. For publishing Harper’s letter, both men were required to vacate their posi- tion in the Club, or face dismissal from the civil ser- vice. According to Lothian (1981: 34), Lloyd had already written a memorandum to Commissioner Harkin in which he called the decision to ship ani- mals from a herd known to be infected to a herd that had never had contact with the disease, “very bad epidemiology,” and “the biologically correct way of dealing with the excess buffalo is to slaughter the excess, thus realizing on the surplus stock.” Professor William Rowan of the University of Alberta visited the Park in 1925 and collected two specimens of Wood Buffalo under permit. He pub- lished an account of his trip in “Country Life.” A typescript of part two of his article is included in the salvaged files. In his final paragraph he wrote “I need not discuss the pros and cons of the proposal here, suffice it to say that it called forth, naturally enough, the most vigorous protests from scientific quarters from all over Canada, from the States and even from England.” Rowan was black-listed by the Department of the Interior as a result. Other condemnations appeared in the two most important scientific journals in the world. Dr. Ritchie, in England, published his objections in Nature. Perhaps the most serious condemnation came from Dr. Thomas Barbour, Director of the Museum of Comparatives Zoology at Harvard University,Boston. In a book review published in Science for 25th November 1932, he referred to the transfer of bison as follows: This, one of the most tragic examples of bureaucratic stupidity in all history, was done against the protests of both Canadian and American naturalists who would rather have seen the surplus bison killed. They were known to be infected with bovine tuberculosis and they are certain to interbreed as well as infect the wood bison, which is a far finer animal and one of great zoological interest because in some respects it seems more like the European wisent than the common American Bison. 3. The Aftermath Wainwright Transplanting a reasonably large number of Bison each year did not solve the overpopulation problem in Buffalo National Park. As early as the winter of 1926-27 more or less regular slaughters were resumed. There is an accounting of the number of Bison killed, and a meticulous record of the money received for the sale of Bison products, but there is no mention of the prevalence of any disease or para- site in the records I have. Table 4 lists the years, and the numbers of Bison killed. Dr. Seymour Hadwen attended many, but not all of the slaughters. The data for Figure 2 come from Hadwen (1942). Numbers killed are shown on the bars. They are Hadwen’s numbers and some of them differ slightly from the government numbers in Table 4. Hadwen suggested that the prevalence may have declined over time. An alternative explanation is that TABLE 4. Number of Bison Slaughtered at Wainwright After Transfers Began. Year 1926- 1927- 1929- 1930- 1931- 1927 1928 1930 1931 1932 Number 2001 1000 525 47 1534 1932- 1933- 1934- 1936- 1937- 1933 1934 1935 1937 1938 1939 ~—s- Total 1220 2000 L000 1522 3246 2918 17013 154 Hi 1012 PER CENT POSITIVE 0.00/= 23 24 25 26 27 THE CANADIAN FIELD-NATURALIST Vol. 116 31 32 33 34 35 36 37 3 YEARS AFTER 1900 fe POSITIVE FOR T.B. FIGURE 2. Prevalence of tuberculosis in bison at Wainwright (After Hadwen (1942). Numbers over bars are sample sizes. the early slaughters concentrated on the oldest ani- mals, which were most likely to be positive for tuberculosis (Table 2). We know that was true for the sample in 1923. It was probably true for 1928 and 1930, because all three of those years had a prevalency rate greater than 70%. If so, as the pro- portion of old animals in the slaughters declined, so would the prevalence of tuberculosis. Wood Buffalo National Park Population: The Wainwright bison were unloaded either at the Hay Camp, located on the west bank of the Slave River about 50 km upstream from Fort Fitzgerald, or at Buffalo Landing, which was about midway between Hay Camp and Fort Fitzgerald. Some accounts mention La Butte as a destination, but it is on the east bank of the river, which is not in WBNP. Park employees at the time of the introduc- tion of the Plains Bison noted that the residents and the newcomers mingled almost at once. That does not mean, however, that there was a great deal of breeding, either with each other, or with the resident Wood Bison, in the year of their arrival. Two thirds of the total were yearlings, and less than 5% of female bison breed as yearlings (Fuller 1962: 21). The first attempt to make a census of the Bison in WBNP was made in 1931 by the Royal Canadian Air Force. They set out to cover the entire winter range and take photographs of all herds of Bison. Something went wrong because they photographed fewer than 2 400 Bison. J. Dewey Soper was sent to WBNP in 1932, where he stayed until 1934 studying the fauna, espe- cially the Bison. He was limited to surface travel — canoe and pack horses in summer; dog sled and snowshoes in winter. Soper did his best to estimate what the numbers might be. His best guess was 12 000, which is within the realm of reality. In 1947, Mr. E. G. Oldham, Superintendent of Forests and Wildlife, attempted an aerial survey. Instead of attempting full coverage he flew strips so placed that he covered one third of the area of the park. He counted 2494 bison so his estimate was 7482 in the park as a whole. Details of his census were in a file at Fort Smith when I arrived there in 1947. In 1949, I made an aerial strip count based on 25% coverage (Fuller 1950). My estimate was not less than 10 000 or more than 12 500. In a repeat aerial census in 1951 (unpublished) I got essentially the same result, which suggests that the population was not growing, but staying more or less constant. All of that came to an end after 1970. In 1969 the flow of water in the Peace River was cut off near its source when the gates were closed on a new dam and the water was held back to fill a reser- voir. The flow was “restored” after the reservoir was filled. That is to say, about the same amount of water comes down the river in a year, but the seasonal dis- tribution of the flow is altered. Spring floods are essentially a thing of the past. Without spring floods every few years the Peace Delta has undergone radi- _ cal change. Perched lakes go dry; what used to be navigable small streams running into Lake Clair are choked with vegetation; sedges are being replaced by grass, and grass by shrubs and trees. All of this has had an enormous effect on the number of Bison in the part of WBNP south of Peace River (Figure 3). 2002 FULLER: CANADA AND THE BUFFALO 155 9.000 8.500 8.000 7.500 6.500 LN CENSUS NUMBER 6.0001 |e 5.500 7.000 ~ 123456 9 1011121 3 14.15 16 17 18 19 20 21 22 23 24 25 26 27 28 YEARS AFTER 1970 LN (NUMBER) —* REGRESSION FIGURE 3. Natural logarithms of counts of bison south of Peace River beginning in 1971. The straight line is the Calculated Regression Line. The equation for the regression line for the log, transformed numbers is: LN(Nt) = 9.141 — 0.086t, with an 7 of 0.887. The rate of decline over 28 years has been 8.6% per year, and the probability that the relationship could have occurred by chance is less than 0.001. Figure 4 shows what has happened to the popula- tion as a whole. The striking similarity between these two Figures shows that changes in the delta have affected not only the southern part of the park but the park as a whole. The equation for this log, transformed regression is LN(Nt) = 9.137-0.051t, with an r of 0.917. The overall rate of decline has been 5.1% per year. The probability that the relationship was due to chance is less than 0.001. North of Peace River the population fluctuated up and down without any visible pattern. The regression line, which actually has a positive slope of 0.5%, is meaningless, with an r? < 0.001. Tuberculosis: Col. Richards ended his summary of correspondence concerning the transfer with two short, but interesting, paragraphs. The information available lends support to the view that as only young animals were being transferred to the Wood Buffalo Park, improvement in their condition might be looked for. It is doubtful whether any of our park officials are in a position to definitely determine to what extent the animals in Wood Buffalo Park are affected with tuberculosis. In these circumstances it would seem that facts are not available to justify the conclusion that tuberculosis is prevalent or endangering the buffalo in Wood Buffalo Park. It is the practice to authorize the killing of a number of aged buffalo in Wood Buffalo Park each year, the meat of which is distributed to charitable institutions and to natives and half-breeds in the northern districts. In view of Mr. Soper’s report it would seem that the ser- vices of a veterinarian should be available at the time of the buffalo slaughter to ensure that the meat of healthy animals only is being disposed of for the purposes men- tioned. When IJ arrived in Fort Smith in 1947 the term was ‘buffalo hunt’ not slaughter. The hunt began in the fall after it got cold enough to preserve the meat. Old males were the targets even though it had been known since the 1923 slaughter at Wainwright that the prevalence of tuberculosis was highest in aged animals, especially males. Park staff did all the shooting. Skinning, butchering, and transporting were the responsibility of the recipients. The Fort Smith hunt took place on the Salt Plains, which could be reached by 4-wheel drive vehicles. The Fort Chipewyan hunt took place on part of the Peace- Athabasca Delta where the only means of transporta- tion was the dog sled, and the whole crew slept in a two-room cabin. In 1950 a professional butcher and a veterinary meat inspector were brought from Edmonton for the hunt at Fort Smith. I worked with the veterinarian, who showed me where to look for signs of tubercu- losis, and what the signs are. I carried on as “inspec- tor” for the animals killed at Chipewyan. Thus ended about 25 years of feeding uninspected carcasses to school children and people on welfare. 156 THE CANADIAN FIELD-NATURALIST Vol. 116 LN CENSUS NUMBER 123 45 6 7 8 9 1011 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 NUMBER OF YEARS (+1970) LN(NUMBER) —* REGRESSION FiGuRE 4. Natural logarithms of counts of bison in Wood Buffalo National Park as a whole beginning in 1971. The straight line is the Calculated Regression Line. For two or three years a mobile “abattoir” was used. It consisted of three buildings on sleds pulled by a tractor. Finally, permanent abattoirs were con- structed at Hay Camp and on the Peace Delta close to Sweetgrass River. A Veterinarian was present at all slaughters once the abattoirs were in use. As an example of the findings, Figure 5 shows the prevalence of tuberculosis in 527 males and 981 females examined at Hay Camp in the years 1952-56 (Fuller 1962: 29). Age Class 0 means calf, Classes 1 to 4 are ages | year to 4 years, Class 5 is Young Adult, Class 6 is Prime Adult, and Class 7 is Old Age. Characteristics used to define each age class are set out in Fuller (1959). It is clear from Figure 5 that the prevalence of tuberculosis increased with age as it did at Wainwright. Almost three-quarters of adult and old males were positive. Overall, 38 per cent of males and 40 per cent of females showed evidence of tuberculosis. There can be no denying that tuberculo- sis had been introduced to Bison in WBNP. Brucellosis (Contagious Abortion): The origin of Brucellosis in the park is a mystery, but it is most likely to be the Wainwright animals. Brucellosis was first confirmed in the park in 1956, which was the year that I left Ft. Smith. Some symptoms character- istic of Brucellosis had been seen during slaughters, but laboratory confirmation was lacking because it had not been possible to keep samples from freezing in the camps and on their way to the lab. The preva- lence of Brucellosis at the Sweetgrass abattoir during its period of use (1957-1958 to 1973-1974) was 39.5%. A total of 1681 individuals were sampled of which 664 were positive. The incidence of tuberculosis at Hay Camp during the same period was 31%. Given these two results, the probability that an animal chosen randomly did not have at least one of the two diseases was 0.417 or about 4 chances out of 10. Recent studies of Bison diseases were reported by Dr. Stacey Tessaro (1989) and Tessaro et al. (1990). Tessaro (1989:5) found “...brucellosis in 18 (25%), and tuberculosis in 15 (21%), of 72 Bison that had died as result of hunting activity, wolf predation, nat- ural accidents, or disease. The combined prevalence of the two diseases in the sample was 42%. Of the 56 Bison that were killed, butchered and utilized by native hunters, 24 (43%) had one or both of these two zoonotic diseases. Three or four Bison seen killed by wolves were severely debilitated by gener- alized tuberculosis, and I suspect that a significant portion of the adult Bison mortality attributed to Wolves is predisposed by disease. Another cow was found to have died of pulmonary tuberculosis. Brucellar arthritis in three Bison resulted in crip- pling, emaciation and death of these animals. Eight other severely lame and emaciated bison were seen in the park, but these animals could not be collected for scientific evaluation.” Tessaro also expressed 2002 FULLER: CANADA AND THE BUFFALO 57 PER CENT POSITIVE AGE CLASS MALES FEMALES FIGURE 5. Prevalence of Tuberculosis by sex and age group in 1508 bison examined at Hay Camp, WBNP, in the years 1952-1956. (After Fuller, 1962). Numbers above bars are sample sizes. concern that the diseases could be carried to the Mackenzie Bison Sanctuary by one or more roaming park bison. Mackenzie Bison Sanctuary In 1958, Dr. N. S. Novakowski located a group of about 200 bison with many of the characteristics of Wood Bison near the northern boundary of the park. In the next few years many of them were trapped and moved to corrals set up on the Salt Plains near Fort Smith. There they were tested repeatedly for tubercu- losis and brucellosis. Reactors were eliminated after each test. Drs. Banfield and Novakowski (1960) examined skeletal material as well as characteristics of the living animals, and declared that the bones and the live animals belonged to the Wood Bison sub- species (Bison bison athabascae). Their opinion has been both questioned and supported several times since they published the results of their study. This is not the place in which to debate taxonomy, but there is no question that those animals and their descen- dants are the closest we will ever see to the original Wood Bison. In 1963, the captives at Ft. Smith were deemed disease-free. Eighteen of them were moved to Ft. Providence where they were released in an area north of the Mackenzie River. The area set aside for them is known as the Mackenzie Bison Sanctuary (MBS). The remainder were sent to Elk Island National Park. For more information about their fate see C. C. Gates et al.(2001). The little group translocated to the MBS now numbers in the region of 2600 head (N.S. Novakowski, personal communication). No road runs through the MBS, but the highway between Yellowknife and Fort Providence runs along its western boundary and travellers frequently see Bison on or beside the road. Although the Mackenzie River separates the MBS herd from WBNP, the river is not an impenetrable barrier. It can easily be crossed on the ice once it has frozen over. The probability of a diseased animal from Wood Buffalo National Park wandering as far as the MBS is low but non-zero. The herd, however, is too valuable to be exposed to the slightest danger if it can be prevented. Conclusions My first objective in this paper has been to show that, clearly, it can be argued that the Canadian gov- ernment made a serious mistake when it gave in to public pressure some 75 years ago. Buffalo National Park was bursting at the seams with Bison. Not only was the range threatened by overpopulation, but a significantly high prevalence of tuberculosis had been demonstrated in 1923 by Hadwen and by the Waddy Report in 1924. The government of the time had at least three choices. One was to continue to slaughter large numbers every year in order to keep the population within the carrying capacity of the 158 THE CANADIAN FIELD-NATURALIST range. The second was to move surplus animals to the new WBNP and take a chance that the diseases would be left behind, and that at least some of the Wood Bison would avoid contact with the intro- duced Plains Bison. The third choice, which was proposed by Hadwen, and backed up by Dr. Clarke, was to slaughter the entire population, allow the range to recover for a few years, and restock Buffalo National Park with healthy animals that were already available in Elk Island National Park. The third option was the only one that guaranteed that no dis- eases would be introduced to the Wood Bison, nor would any hybridization between Plains and Wood Bison take place. Hewitt’s suggestion to sell surplus Bison to farm- ers faced several difficulties. Among the most seri- ous was that the Park lacked the facilities for holding and testing hundreds of animals every year in spite of Graham’s repeated pleading for the necessary equipment. It would have been highly unethical, and probably illegal, to knowingly sell diseased animals that might end up on the farmer’s dinner table. The government put its faith in the second option, which rested on the three untested and unproven hypotheses set out in this paper. The test of those hypotheses was the transfer itself and, as we have seen, all three hypotheses were untrue. Unfor- tunately, the experiment was non-reversible. My second objective in preparing this paper was to discharge the obligation I assumed regarding Maxwell Graham. I believe that the excerpts from the old files show that Graham was devoted to the Bison at Wainwright. He showed continuous con- cern for their health and was devastated when he learned that tuberculosis was present in the herd. He was Cleared of all blame by two of his superiors in the civil service, one of whom was the Deputy Minister. He played no role in the decision to trans- fer young bison to the newly formed Wood Buffalo National park, and he had left the National Parks Branch some two years before the transfer began. His one transgression was the one-page letter he had published in The Canadian Field Naturalist that sup- ported the three failed hypotheses. We will probably never know whether he wrote that letter on his own initiative or whether he was “asked” by one of his superiors to write it. There is no doubt that the annu- al calf crop in the mid 1920s at Wainwright faced an uncertain future, and I can understand why Graham sincerely believed that the transfer gave the young animals their best chance for survival. In August of 1990 the Report of the Environmental Panel on Northern Diseased Bison was made public. The recommendation of the Panel, of which I was a member, was that all the Bison in Wood Buffalo National Park should be slaughtered. During and after the slaughter, Bison from Elk Island National Park should be bred in small enclo- Vol. 116 sures in the north so that restocking with disease-free animals could take place as early as possible. The recommendation met with opposition on many fronts and, shades of 1923, the government backed down once again. The government decided to “study” the situation. Ten years on, no action has been taken. So here we are in the new Millennium, and we are faced once again with a situation similar in many respects to that faced by the managers of Buffalo National Park in 1923-24. There is a herd of bison with a high prevalence of two infectious diseases and a declining population. There is another herd that is disease-free and flourishing. There is a low, but non-zero probability of infection of the clean herd by diseased animals wandering from their prop- er home. The Panel recommended total slaughter and replacement, but it did not recommend slaughter just for the sake of killing. We must keep in mind what happened at Wain- wright when the government refused to do a com- plete slaughter after it was shown that up to 75% of old animals were infected with tuberculosis. In 1923, the Bison population was under 7000. The total loss of life would have been less than 7000 if the whole herd had been slaughtered as Dr. Hadwen recom- mended. Ultimately, the protesters and a weak gov- ernment were responsible for the deaths of 17 000 Wainwright Bison (Table 4), for introduction of dis- eases to the pristine population of Wood Bison, and for contaminating the majestic Wood Bison by hybridization with Plains Bison. The situation in WBNP, unlike that in Wainwright, has actually improved with time. In 1990, the Bison population was more than 3300. In 1998, it was just over 2300 and declining. Some of the annual slaugh- ters at Wainwright were as large as, or larger than, what would be needed for a final clean-up in WBNP. There is no logical reason for further delay. Literature Cited Allen, J. A. 1875. Annual Report of U.S. Geological and Geographical Survey. Part 3: 443-588. Banfield, A. W. F., and N. S. Novakowski. 1960. The survival of the Wood Bison (Bison bison athabascae Rhoads) in the Northwest Territories. National Museum of Canada, Natural History Paper 8. Cameron, A. E. 1923. Notes on buffalo: Anatomy, patho- logical conditions, and parasites. The Veterinary Journal 79: 331-336. Cameron, A. E. 1924. Some further notes on buffalo. The Veterinary Journal 80: 413-417. Environmental Panel. 1990. Northern Diseased Bison. Federal Environmental Assessment Review Office. Foster, J. E. 1992. The Métis and the end of the Plains Buffalo in Alberta. Pages 61-77 in Buffalo. Edited by John Foster, Dick Harrison, and I. S. MacLaren. Univer- sity of Alberta Press. Franklin, J. 1823. Narrative of a journey to the shores of the polar sea in the years 1819, 20, 21, and 22. Facsimile reprint. M. G. Hurtig Ltd. Edmonton, 1969. 768 pages. 2002 FULLER: CANADA AND THE BUFFALO 159 Fuller, W. A. 1950. Aerial Census of Northern Bison in Wood Buffalo Park and Vicinity. Journal of Wildlife Management. 14 (4): 45-51. Fuller, W. A. 1959. The horns and teeth as indicators of age in bison. Journal of Wildlife Management 23: 342-344. Fuller, W. A. 1962. The biology and management of the bison of Wood Buffalo National Park. Canadian Wildlife Service, Wildlife Management Bulletin Series 1, Number 16. Gates, C. C., R. O. Stephenson, H. W. Reynolds, C. G. vanZyll de Jong, H. Schwantje, M. Hoefs, J. Nishi, N. Cool, J. Chisholm, A. James, and B. Koonz. 2001. National Recovery Plan for the Wood Bison (Bison bison athabascae). National Recovery Plan Number 21. Recovery of Nationally Endangered Wildlife (RENEW). Ottawa, Ontario. 50 pages. Graham, M. 1924. Observations in the Wood Buffalo Park 1922. Canadian Field-Naturalist 38: 189. Hadwen, S. 1942. Tuberculosis in the buffalo. Journal of the American Veterinary Association. 100: 19-22. Harper, F. 1925. Letter to Editor. Canadian Field- Naturalist 39: 45. Hearne, Samuel. 1795. A journey from Prince of Wales’s Fort in Hudson’s Bay to the Northern Ocean. Undertaken by Order of the Hudson’s Bay Company for the Discovery of Copper Mines, a Northwest Passage, &c. In the Years 1769, 1770, 1771, & 1772. Facsimile reprint of the first edition. Edmonton, Alberta: M.G. Hurtig Ltd., 1971. 458 pages. Hewitt, C. G. 1919. The coming back of the bison. Natural History. 19: 553-565. Hewitt, C. G. 1921. The conservation of the wild life of Canada. Charles Scribner’s Sons. 344 pages. Hornaday W. T. 1889. The extermination of the American bison, with a sketch of its discovery and life history. Annual Report of the Smithsonian Institution, 1887, Volume II (Washington DC: Government Printing Office, 1889.) Howell, A. B. 1924. Letter to Editor. Canadian Field- Naturalist. 38: 118. Isenberg, A. C. 2000. The destruction of the bison: an environmental history, 1750-1920. Cambridge University Press. 206 pages Lothian, W. F. 1981. A history of Canada’s National Parks Volume 4, Pages 25-32. Lumsden, H. G. 1984a. A tribute to Charles Henry Douglas Clarke, 1909-1981. Canadian Field—Naturalist 98: 379-384. Lumsden, H. G. 1984b. Bibliography of C. H. D. Clarke, Canadian Field-Naturalist 98: 385-391. Mackenzie, A. 1801. Voyages from Montreal on the River St. Laurence [sic], through the continent of North America, to the frozen and Pacific Oceans; In the years 1789 and 1793. London and Edinburgh. Mitchell, J. A., and C. C. Gates. 2002. Status of the Wood Bison (Bison bison athabascae) in Alberta. Alberta Wildlife Status Report No.38. Alberta Sustainable Resource Development. 34 pages. Pike, W. 1892. The barren ground of northern Canada. Macmillan and Co. London. 300 pages. Polziehn, R. O., C. Strobeck, J. Sheraton, and R. Beech. 1995. Bovine mtDNA discovered in North American bison populations. Conservation Biology 9: 1638-1643. Preble, E. A. 1908. A biological investigation of the Athabaska-Mackenzie Region. North American Fauna (27). 574 pages. Rowan, W. 1929. Canada’s buffalo. Country Life. 348-340. Roe, F. G. 1951. The North American Buffalo: A Critical Study of the Species in Its Wild State. University of Toronto Press. 957 pages. Russell, F. 1898. Explorations in the Far North. Univer- sity of Iowa. 290 pages. Saunders, W. E. 1925. Letter to Editor. Canadian Field- Naturalist 39: 118. Seton, E. T. 1911. The Arctic Prairies: A canoe journey of 2,000 miles in search of the caribou being an account of a voyage to the region north of Aylmer Lake. Inter- national University Press. New York. 308 pages. Soper, J.D. 1939. History, range and home life of the northern bison. Ecological Monographs. 11: 347-412. Tessaro, S. V. 1989. Saving the wood bison in its tradi- tional range: The threat of tuberculosis and brucellosis. Wildlife Veterinary Report. 2: 5—6 Tessaro, S. V., L. B. Forbes, and C. Turcotte. 1990. A survey of brucellosis and tuberculosis in bison in and around wood buffalo National Park, Canada. Canadian Veterinary Journal. 31: 174-180. Accepted 12 April 2001 Received 11 March 2002 Book Reviews ZOOLOGY Birds, Mammals & Reptiles of the Galapagos Islands By Andy Swash and Rob Still. 2000. Pica Press and WildGuides, Yale University Press, New Haven and London. 168 pp., illus. U.S. $24.95. I was delighted to find a sturdy, handy-sized guide to all the animals of the Galapagos Islands. For the traveler, size as well as quality is important. This book certainly is the right size and packs all the essential information into one volume. It covers all of the resident mammals, reptiles, and birds. It also covers the migrant and vagrant birds as well. There are introductory sections on geography, climate and habitat. So the size and content are good, but what of the quality of the information? This is somewhat of a mixed bag. The introductory sections are logical, concise, and clear. They contain as much as I need to have in a book I plan to carry in my pocket while trudging across a tropical island. The bird section begins with an introduction to the major bird groups. The authors have used a modified taxonomic group- ing that works well. This leads into the main section on the bird species. This book relies on computer-manipulated digital images as the main form of illustration. As a result the plates look more like a “standard” field guide of the Peterson type. This is an improvement over the old sequence of small pictures. However, my arguments against photographs in field identifications still hold. For example, the Black Petrel and the Hoary Bat have a strong rosy tinge, giving a distorted impression of their true colours. The juvenile Yellow Warbler has a powder blue head. Several other birds had odd colours, including lilac-toned Lava Gull and Sora. The strangest transformation is on the female Beited Kingfisher. The breast bar is clearly visible, but the belly bar is so obscure a novice might miss it. Yet more strange is the breast bar is distinctly chestnut, instead of gray. Is this computer manipulation gone awry? Other birds on the same page as the odd ones were normally coloured. I also wonder why the authors did not change the backgrounds to give more contrast with the birds. They have brownish herons on a brownish background; grayish mockingbirds on a grayish background, and so on. If they wanted to manipulate why not follow their own fine example, the flycatcher page. This shows the birds clearly and in good proportion. The text explains that the light morph of the Wedge-tailed Shearwater is more likely yet the plate depicts the dark morph. I would also prefer that the authors had emphasized the flight of oceanodrama petrels, a much more useful characteristic in the field. One plate is devoted to the heads and beaks of Darwin Finches. The heads are depicted against a graph paper background and the outline of each bill is picked out in an unobtrusive gold line. Coupled with the description this is a useful tool for the inex- perienced visitor. The whale section is, for the most part, well done, with the species being depicted as you see them from a ship. While this does not show much of each species, it does present a realistic view of a typical field sighting. It also demonstrates the difficulties of field identification. I was not happy with the photo- graph of Cuvier’s Beaked Whale. It was dark and showed little of this species’ characteristics. The authors did mention the possible spotting on a Sei Whale. The Blainville’s Beaked Whale is coloured red. I have not seen a Blainville’s Beaked Whale, but the other beaked whales I have seen were at best described as brownish. I would be surprised to find a maple leaf red specimen. I would like to see the authors add information on seasonal distribution. I realize most of the birds are year round residents, but there are others I question. Are the “resident” Waved A present all year or they disperse at the end of their breeding season? This information would also be valuable with scarce migrants. When can we expect to see skuas or jaegers? Despite my comments, this is a useful field book. It is unlikely that anyone would confuse any of the species, despite the odd errors the species that could be confused are well covered. The book covers all the Galapagos animals in one handy portable vol- ume. So I will try it in the field in late October this year. In the meantime I will be using other higher quality, but far less portable, texts for research. Roy JOHN _ 2193 Emard Crescent, Beacon Hill North, Gloucester, Ontario K1J 6K5 Canada 160 2002 BOOK REVIEWS 161 Wisconsin Fishes 2000: Status and Distribution By John Lyons, Philip A. Cochran, and Don Fago. University of Wisconsin Sea Grant Publication No.WISCU-B-00-001. University of Wisconsin Sea Grant Institute, 1975 Willow Drive, Second Floor, Madison, Wisconsin 53706-117. e-mail: linda@sea- grant.wisc.edu. 87 pp. U.S. $10. One wonders if the Electronic Age will make peer book reviews less important, or more important. An eight-page description of this book is available at http://www.seagrant.wisc.edu/greatlakesfish/Lyons.h tmf. This is my first experience evaluating a publica- tion with most of the routine items, or samples, read- ily at hand. The header on each of the eight pages of the Internet description uses the statement “Wisconsin Fishes 2000/Fish of the Great Lakes by Wisconsin Sea Grant.” This Great Lakes emphasis is not men- tioned in the book. Great emphasis is placed, by the authors, on the literature since Becker’s (1983) Fishes of Wisconsin, and the fact that this publica- tion updates the information on “occurrence, taxo- nomic status, and abundance of fishes in Wisconsin’. Fishes in the waters of the Wisconsin portions, and at times the adjacent Michigan portions, of lakes Michigan and Superior are, however, included. Individuals interested in the status and distribution of fishes in Wisconsin, and comparisons with those in the geographic areas around Wisconsin in the USA and Canada, owe a debt of thanks for the extensive and constantly updated information since Becker’s 1983 book, in the 22 related publications by these authors and coauthors, and those of several other authors. Much of the information in this book was derived from the “Master Fish File”, a database including 22 000 Wisconsin fish collections from 1900 to 1999. The database can be accessed through the web site of the Wisconsin Department of Natural Resources (www.dnr.state.wi.us). This book includes sections entitled Summary, Introduction, Materials and Methods, Overview of Changes in the Wisconsin Fish Fauna, Species Accounts, References, Index to Common Name, Index to Species by Scientific Name, and eight colour plates. The overview of changes includes a table with sections entitled Native Species (147), Established Non-Native Species (14), and Transient Non-Native Species (19). In total those sections include 180 species. One of the problems of lists is the neces- sary, sometimes arbitrary, decisions required when authors decide where to put various species. Readers may not have made the same decisions. Five species not included in earlier publications are included here. Those are Kokanee, Oncorhynchus nerka, Threespine Stickleback, Gasterosteus aculea- tus, White Perch, Morone americana, and Round Goby Neogobius melanostomus. There are species accounts for 182 forms, ranging in length from a few lines to several pages. The emphasis, in length, is on the more recently discov- ered species, native and introduced. Many of those were not given full treatment in Becker’s 1983 book. Only eight of the species accounts include a “spot” map of those locations at which the species has been collected, or observed by one of the authors. The longer species accounts usually include Common Name; Scientific Name; Description, including means of separation from look-alike species; Systematic Notes; Distribution, Status, and Habitat; Biology; and Importance and Management. The dis- tribution statements, and absence of a map, in the shorter Species Accounts leaves the reader unable to determine if the species occurs in lakes Michigan and/or Superior. Some species accounts deal with extirpated species, and species rediscovered as pre- sent. The book includes eight colour plates tipped into the front before the title page, somewhat like a Frontispiece. The plates are of seven recently dis- covered forms. There is one unfortunate problem. The caption of one of the two plates for the Round Goby, Neogobius melanostomus, incorrectly identi- fies the “fused pectoral fins”, whereas the photo clearly indicates that it is the pelvic fins which are fused forming the diagnostic “sucking disc” of gob- ies. There are a number of interesting points to be derived from the lists and accounts. The abundance of natural reproduction of Lake Trout, Salvelinus namaycush in the Wisconsin portion of Lake Superior has led to stocking being discontinued there. The same is apparently true for Chinook and Coho salmon, Oncorhynchus tshawytscha and O. kisutch. In contrast, for Pink Salmon, O. gorbuscha, there has been no natural spawning in Lake Superior streams since the late 1970s and the species is now rarely seen in Wisconsin waters of lakes Superior and Michigan. There have apparently been no records of the European Flatfish, Platichthys flesus from Wisconsin waters of lakes Michigan or Superior. This is in contrast to the fact that Ontario records for Lake Superior are almost as numerous as those for Lake Erie. The Warmouth, Lepomis gulo- sus has now been reported from (the southern part of ?) the Lake Superior Basin. This species is easily confused with the Rockbass, Ambloplites rupestris, and even though Lake Superior may be more of a barrier (cold) to immigration, with the experience on Lake Erie its arrival in Lake Superior should be watched for. There is, presently, a great emphasis on mainte- nance of biodiversity, rehabilitation of native species, and the controversy around “Is more bet- ter?”. This commitment cannot hope to succeed 162 without constant surveys, species status reports, checklists, and publications like this one and the oth- ers which have appeared in Wisconsin. Too often, surveys and updates, such as this one, are deemed less important than “more sophisticated” research. I hope the people in Wisconsin will continue to be supported philosophically and financially to set an example for other Political areas. I found the book interesting and useful. I’m certain that fish biologists in Wisconsin are finding it useful, as will the next person charged with updating a checklist of the fish- es of the Great Lakes Basin. THE CANADIAN FIELD-NATURALIST Vol. 116 References Becker, G. C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison, Wisconsin. Robins, C. R.,R. M. Bailey, C. E. Bond, Jr., Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott. 1991. Common and sci- entific names of fishes from the United States and Canada. Fifth Edition, American Fisheries Society Special Publication 20, Bethesda, Maryland. E. J. CROSSMAN Curator Emeritus, Ichthyology, Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario M5S 2C6 Canada Marine Mammals of the Pacific Northwest: A Concise and Comprehensive Waterproof Guide By P.A. Folkens. 2001. Harbour Publishing, Madeira Park, British Columbia. 8 pp., illus. $9.95. The increasing development of the whale-watch- ing industry has resulted in the demand and creation of books and identification guides for sea mammals, seabirds, and other parts of the marine ecosystem. In this regard, and looking back on a history of whale watching for over 90 years, the coast of British Columbia has received most of the publication activ- ity in Canada. Despite its catchy title, this guide by Pieter Arend Folkens is more a leaflet than a guide. It consists of three text pages and five pages of drawings and pho- tos full of information for the whale watcher in the field. Although the leaflet is printed on “waterproof, UV resistant synthetic film made from a 100% recy- clable, environmentally inert material containing no forest products (similar to milk jugs)”, the user might actually have difficulties using it on an off- shore whale watching trip during periods of strong wind or high waves; the light leaflet could easily fly away and the small print is hard to read when on a rolling boat. However, the compressed text gives a nice summary and overview on 31 sea mammal species in the area; it even mentions Steller’s Sea Cow which was hunted to extinction by 1768. In addition, major whale watching locations in British Columbia and Alaska are named; but none are specifically reported for Oregon and Washington (as the title would imply). Most of the eight pages of the field guide are devot- ed to drawings and to fine pictures from the author and several others. The reader might find the distinction between Mysticetes (Baleen Whales), Odontocetes (Toothed Whales), and Small Cetaceans a little unclear from the arrangements of the drawings. Very helpful . and informative is the page about “Common visible behaviors and terms” allowing to link sea mammal sightings to a classified set of behaviour types. Helpful also is that images of fluke displays are presented for species that are known to show such behaviour. Even the body sizes of newborn sea mammals are given. All measurements are made in SI units, and the conversion factor for feet is provided. Of interest to the general audience might be the section “Marine Mammal Watching Guideline,” also presented on the web [http://www.fakr.noaa.gov/ protectedresources/mmviewingguide.html] (Note that the old URL www.nmfs.gov/prot_res.html and given in the guide was updated). In addition, contact addresses and a web address (revised to http://www.fakr.noaa.gov/protectedresources/strand- ings.htm) are given for sightings of stranded sea mammals (Canada: Department of Fisheries and Oceans 800 465-4336; or the Whale Reporting & Stranding Line 800 665-5939). As in many other field guides, the text suggests some field marks and details for species identifica- tion and separation that normal whale watchers might not be able to apply, or which are not really realistic. For instance, Sei Whales are supposed to be differentiated from Fin Whales by a fin angle of over 45 degrees; male Beaked Whales (Genus Meso- plodon) ideally can be identified by the location of teeth and jaw line (which is, for most of the time, covered by the ocean); phocids (true seals) differ from otariids (Sea Lions and Fur Seals) by their hair and small nails on their foreflippers. Overall, fea- tures like these might be very hard to recognize for the untrained as well as for the trained observer, par- ticularly when observations are made of moving ani- mals some distance from a shaky boat, with binocu- lars. Rather than focusing on classical small-scale features, an outline of the use of proportions and shapes could be more useful for distinguishing species. Counterproductive for a field guide might be the point that Beaked Whales, the species group that lacks most knowledge on distribution and where whale watchers could indeed contribute greatly to science, are described as the “most difficult whales to identify correct.” No further help or details are given for the interested whale watcher. 2002 BOOK REVIEWS 163 For my taste, the “Habitat and Symbol Keys” that are supposed “to narrow possibilities in a particular era” and link sea mammals with “habitat” are not really helpful. Many whales migrate across habitats and the regular observer has no real way to tell “tem- perate” habitats apart from “cool temperate” ones. The meaning of the orange W habitat class presented for the False Killer Whale will likely remain a mys- tery to the reader because its meaning is unexplained in the guide. For pinnipeds, their “calls” and the mention of rookeries could have been helpful. Manitoba Birds By Andy Bezener and Ken de Smet. 2000. Lone Pine Publishing, Edmonton, Alberta. 176 pp., illus. $17.95. I often wonder in which market book writers think they will sell their product. My first reaction to this compact little book, covering only 145 species (out of the official list of 406) was that it was too simple for most birdwatchers. And it probably is. Then I realized it was ideal for scout leaders. It gives a short, easy-to-read account of a bird’s character, and a sentence or two on identification, size, status, habi- tat, nesting, feeding, voice, similar species, and the best sites in the province (to see the bird). An illus- tration and a range map accompany this information. This is just about the right level of detail for a scout leader teaching his troop about birds. It is also good for schoolteachers, new birders, and children. While it is intended for use in Manitoba, it would also be a useful guide throughout the prairies. The introduction has a map of the natural regions showing the best 50 birding sites. A dozen sites have a short write-up that includes the most prominent bird species. I liked this addition to the text. I thought beginners and visitors would be able to use it as a starting point when organizing trips. Each account has a small illustration. While most of these are reduced (and reversed) versions from the main text, five species are different and not covered in the core section of the book. I counted 145 in the main text, so there is a total 150 species illustrated. These additional species, which this guide explains you can find most easily in the places mentioned, would be better included in the central body of the book (in replacement for some of the rarer or more difficult to see birds). The authors claim the birds selected are the most common and easily found within the province. I question some of the choices. They have chosen, for Overall, it appears that this “guide” is an excerpt of a better and larger guide book from the same author. It is useable in the field, but does not replace the real and classical guide books. FALK HUETTMANN Centre for Wildlife Ecology, Biology Department, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia V5A 1S6 Canada. example, Burrowing Owl, a rare nester in the south west corner of the province and Screech Owl which nests along a narrow strip close to the U.S border (the range map for this owl seems a trifle optimistic). Yet they left out Hawk Owl, a widespread bird which nests over the northern three-quarters of the province. Birds seen easily in the north (Churchill) but not in the south, such as Bonaparte’s Gull, jaegers, ptarmigan, and plovers, are also not includ- ed. Generally, though, I found the choices to be rea- sonable for the populated south of the province. I would be surprised if House Sparrow is not found at northern towns like Lynn Lake and Thompson (the range map does not suggest this.) The illustrations by Gary Ross, Ted Nordhagen, and Ewa Pluciennik are, for the most part, very good. They generally show a typical adult in summer plumage. For some, but not all, species the differ- ences between male and female are shown. Winter visitors are depicted in the appropriate non-breeding plumage. Many birds are shown in flight too. I was very impressed with the shape, attitude, and techni- cal detail of each species. I did find a few that I thought could be improved (Warbling Vireo is too pale and Olive-sided Flycatcher is not heavily built or boldly coloured enough), but overall the quality is first rate. This is a compact (14 X 21 X I cm) little book that will easily slip into a pocket or pack. The plasti- cized cover and high quality production suggests it should last well in the field. A quick colour-coded guide to the bird families on the back cover will also help out novices to find the appropriate text. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Gloucester, Ontario K1J 6K5 Canada SS 164 Heron Conservation Edited by James A. Kushlan and Heinz Hafner. 2000. Academic Press, London and San Diego. xvi+480 pp. illus. This publication is a cooperative one, with 19 major contributors and a multitude of cooperators from the Herons Specialist Group, an international network of experts in the field of herons and wetland protection. Their objective was “to synthesize and summarize the state of knowledge of the conserva- tion needs of herons throughout the world.” The book opens with eight chapters on the status of herons in major geographic regions — either con- tinents or sub-continents. The herons of each area are reviewed species by species, with estimates of their numbers, or at least their status and conserva- tion needs. Of necessity there is much variation in the coverage, with considerable detail available on European species and lamentably little for many Asian and South American ones. Next come seven chapters on various aspects of heron conservation, covering such topics as habitat conservation, aqua- culture, and environmental contaminants, then a chapter on research and information needs, and final- ly a synthesis of the conservation needs for the most vulnerable species. There is an extensive bibliogra- phy and a well-organized index. Any book devoted to the conservation needs of any group of species on a global scale these days makes for depressing reading, and this work is no exception. Information is lacking on even the most basic aspects of life history and distribution of many more elusive herons, and the sad chronicle of habitat loss, exploitation, and persecution is common to all too many species cross the world. Still, this is hardly news to anyone interested in birds, and I found myself wondering about the audi- ence for a work of this kind. Action on any individu- al species or group of species will necessarily be quite specific to the local conditions and the needs of the birds. Much of the information given here, how- ever, is very general, perhaps inevitably so given the enormous scope of the book. One would think this information would already be well known to those working in the field, yet at the same time the very specific scope of the book limits its appeal to a broader audience. In an attempt to assess its relevance to those species that I am most familiar with, I placed partic- ular emphasis on the North American chapter. I was left with a distinct sense of disquiet. Its general tone of optimism runs contrary to my own experience on an admittedly very limited, local scale. It’s true, of course, that our problems are minor when compared to the enormous challenges elsewhere, but at the same time the problems are real. I’m not sure that cheerfully quoting the Canadian government’s poli- cy of “no net loss” of wetlands is particularly helpful THE CANADIAN FIELD-NATURALIST._ Vol. 116 when — as the authors themselves concede — it only applies to the 29% of wetlands that are federal- ly owned, most of which are north of significant heron range. Many of the wetlands I know are pri- vately or provincially owned, and dying the death of a thousand tiny cuts, while supposedly safeguarded by underfunded agencies reporting to unsympathetic governments. I question the statement that “Tt is like- ly that, overall, the loss of wetland has been halted in Canada...” I found myself equally at odds when considering some individual species, again finding more seeming optimism than I can share. On the Least Bittern, for example, the authors do stress the lack of good data and the fact that in 1986 birders were already expressing concerns about the species. But then the 1965-1979 Breeding Bird Surveys, of all things, are cited as indicating “some evidence” of increases. The reference is correct as far as it goes, but when one considers that in 1997, for example, a grand total of 18 Least Bitterns were recorded on only seven of 1832 BBS routes continent-wide, one wonders how useful its’ information is in the present context. But perhaps the authors are indeed right on a con- tinental scale, and then the bothersome part is their conclusions could easily seem to justify complacen- cy generally. They would doubtless respond that all the caveats about lack of data and “alteration of wet- land functions” [replacement of marshes with boat docks, perhaps?] are there. Indeed they are, but they only “temper” the rosy picture they have already drawn, poor data or not. However, none of the other continental authors seem quite so comfortable with the conditions their herons face, and the book as a whole seems to achieve its objective of pinpointing those areas that need action to conserve heron species world-wide, and the kinds of action needed. The chapters on research and information needs, and the final synthe- sis should be particularly useful in this regard. If one issue emerges as of paramount importance every- where as a critical first step, it is the pressing need for more data, and particularly more precise data, to allow species’ status to be understood. This book is an important one for anyone interest- ed in heron conservation. While North American readers may share my irritation at the authors’ approach, careful reading will still yield a reasonable summary of the issues we face in conserving our herons. However, the book’s appeal to a broader audience is necessarily limited. CLIVE E. GOODWIN 1 Queen Street, Suite 401, Cobourg, Ontario K9A 1M8 Canada 2002 BOOK REVIEWS 165 The Field Guide to the Birds of Australia By G. Pizzey and F. Knight. 1999. Revised Edition. Angus & Robertson, Sydney, Australia. 576 pp., illus. AUD $39.95. The success or failure of any field guide depends on its ability to convey information necessary for correct species identification. The latest edition of The Field Guide to the Birds of Australia by veteran author Graham Pizzey and illustrator Frank Knight accomplishes this objective, and does so in an appealing format. The introduction to this edition provides a primer for those new to birding, or for those wishing to “brush up” on their terminology. A generous mix of illustrations, text, and examples are used to familiar- ize the reader with descriptions of anatomy, field marks, and behaviours. Although reading this section is not essential to understand the field guide, it will assist the novice birder in gaining the most from a birding excursion. Following the introduction, there are species accounts for 778 native and introduced species with 700 distribution maps. Typically, there are 3—4 species accounts per double page with text and color illustrations opposite each other. Descriptions for each species include information on (1) other names, (2) similar species, (3) voice, (4) habitat, (5) breed- ing season, (6) nests, (7) eggs, and (8) range and sta- tus. The range information is particularly useful since it refers to place names, easily found on maps conveniently included inside both covers. The colour illustrations are well done and depict male, female, and juvenile plumage; races and moult patterns also are provided to facilitate identification. The species accounts are followed by an introduc- tion to the bird families of Australia. General charac- teristics of each family (i.e., behaviour, food, and range) are presented in the same order in which they appear in the species accounts. The world range of each family is provided, as are the number of species within each family and the number of those that occur in Australia (both native and introduced). There are few flaws in the guide. The species accounts and illustrations are helpful and well orga- nized. It would have been useful, however, to include migratory information on the distribution maps, perhaps with seasonal ranges depicted with separate colours. Distribution maps for some intro- duced or rare species are not provided and would have been a nice feature. As a new resident of Australia, and unfamiliar with the birds I was to encounter, I found this field guide to be of great assistance. It is small and hardy enough to toss into a daypack, but also would make a useful reference for anyone interested in Australian birds. SHAWN MorRRISON 8/1 Totterdell St, Belconnen, Australian Capital Territory, 2617, Australia Katydids and Bush-Crickets: Reproductive Behavior and Evolution of the Tettigoniidae Darryl T. Gwynne. 2001. A volume in the Cornell series in arthropod biology (John Alcock, editor). Cornell University Press, Ithaca. xii + 317 pp., illus. U.S. $42.50. Darwin (1871) introduced the theory of sexual selection as an attempt to explain the evolution of such traits as bright colours, huge feather plumes, antlers, and other male adornments, features that by their highly conspicuous nature, seemed highly unlikely to confer survival advantages on their bear- er. Sexual selection (differential reproductive suc- cess resulting from competition for mates) as a topic of investigation lay essentially dormant for about a century. However, since the mid-1970s, interest in sexual selection has blossomed (yes, it occurs in plants, too!), following the recognition that it is a major evolutionary agent. Sexual selection is the underlying theme of Gwynne’s book, which in this respect is similar to those of Eberhard (1985, 1996), Gould and Gould (1989), and Andersson (1994). However, it has the advantage over these contributions in that it deals with the phenomenon in a specific taxon, the family Tettigoniidae, which has allowed the author to pre- sent his ideas on a solid platform of basic biology. Indeed, the author’s successful aim is not only to interest students of sexual selection, but to show nat- uralists what a fascinating group of insects the tet- tigoniids are. For field naturalists, the early chapters, which deal with evolution, diversity (more than 6000 species arranged in 17 subfamilies), life histories, food preferences, natural enemies, defence mecha- nisms, and sound communication, will probably be of most interest. Behaviorists, especially those who focus on reproductive behaviour, will find much food for thought in the second half of the book. However, animal biologists in general will find the entire volume contains much to hold their attention. The book is profusely illustrated with line draw- ings, black-and-white photographs (some of which, 166 THE CANADIAN FIELD-NATURALIST unfortunately, lack contrast), and four pages of color plates. The text is generally well written and Gwynne’s approach (present the hypothesis, then examine the evidence) is highly recommended as a “model” for developing researchers. Occasionally, the author “overindulges” by giving too many exam- ples to illustrate a point, and the editor might have used a sharper pencil to remove redundancy and rep- etition and a sharper eye to ensure that the indexing is accurate. I stress, however, that these are minor “complaints” and my overall opinion is that Gwynne deserves full credit for his synthesis of the biology of this major, yet not well known, insect group. A Manual for Wildlife Radio Tagging Robert E. Kenward. 2001. Academic Press, San Diego, California. 311 pp., illus. U.S. $65.00. This book is a significant updating of “Wildlife radio tagging: equipment, field techniques and data analysis” (Kenward 1987). The book incorporates much new information developed since the iate 1980s and includes all of the original topics from the earlier volume but is 70% larger. Kenward starts this book with the logical least common denominator, that is, deciding whether radio tagging is an appropriate option for collecting data from a subject animal and planning projects. Once the decision to radio tag is made, you proceed through the chapters of the book to determine the appropriate equipment, how to obtain equipment, how to make and attach radio tags, proper radio tracking technique, and data analysis. It is unfortunate that this volume contains many errors, particularly in the literature cited. Some of the errors can make it difficult for less knowledgeable readers to locate and obtain references. Bird Census Techniques By Colin J. Bibby, Neil J. Burgess, David A. Hill, and Simon H. Mustoe. 2000. 2nd edition. Academic, London. 301 pp., illus. U.S. $55.00. The global concerns for the plight of long distance migratory birds and efforts to develop comprehen- sive conservation plans in North America encom- passing all avian species make the need for statisti- cally valid bird census techniques evident. This book, updating and expanding the first edition, fills that need. The authors have incorporated the vast literature of bird censusing to make this volume a comprehen- sive reference. Each major bird counting technique is Vol. 116 Literature Cited Andersson, M. 1994. Sexual Selection. Princeton University Press, Princeton, New Jersey. Darwin, C. 1871. The descent of man and selection in relation to sex. John Murray, London. Eberhard, W.G. 1985. Sexual selection and animal genitalia. Harvard University Press, Cambridge, Massachusetts. Eberhard, W. G. 1996. Female control: Sexual selection by cryptic female choice. Princeton University Press, Princeton, New Jersey. Gould, J., and C. G. Gould. 1989. Sexual selection. Scientific American Library, New York. CEDRIC GILLOTT Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada This book is aimed at the beginning researcher contemplating use of radio tagging as well as the more experienced. Even with the errors as noted above, because of Kenward’s depth of knowledge and extensive experience with tracking, this book is of definite value to any and all researchers that are considering or applying telemetry to their research. It is highly recommendable for its thoroughness, time- liness, and user friendly organization. Those already owning the earlier volume should still consider pur- chasing this new version. Literature Cited Kenward, R. E. 1987. Wildlife radio tagging: equipment, field techniques and data analysis. Academic Press, San Diego, California. ROGER D. APPLEGATE Research and Survey Office, Kansas Department of Wildlife and Parks, P. O. Box 1525, Emporia, Kansas 66801-1525 USA given a thorough treatment in cook-book fashion that makes it possible for anyone to lay out and conduct the count. The book also provides a general back- ground on design of bird censuses and statistical issues. There is information on use of computer soft- ware provided in the chapter on line transects. The book has chapters on applying census meth- ods to specific species. These chapters deal with species that are largely European. However, in most cases, the techniques can be applied to closely relat- ed North American species. For example: Willow Ptarmigan (Lagopus lagopus) and Red Grouse (L. I. scoticus). 2002 BOOK REVIEWS 167 Bibby et al. is relatively free of errors. Those noted were minor and do not detract from the read- ability or utility of the book. I would like to have seen some thoughtful discussion of issues relating to the use of index methods as opposed to methods that provide an estimate. However, overall this book has met the authors’ objectives of providing a compre- hensive reference on bird censusing. I recommend this book to all who are conducting or will be con- ducting such field studies. For those owning the first edition, I recommend purchasing this second edition. ROGER D. APPLEGATE Research and Survey Office, Kansas Department of Wildlife and Parks, P. O. Box 1525, Emporia, KS 66801- 1525 USA Flying Foxes: Fruit and Blossom Bats of Australia By L. Hall and G. Richards. 2000. Krieger Publishing Company, Malabar, Florida. viii + 133 pp., illus. Cloth U.S. $29.50; paper U.S. $21.50. I have had a great interest in flying foxes since I first saw them roosting in a tree in Indonesia about 20 years ago. Their size and open nesting sites in an urban area made them far more spectacular than the Canadian cave species that I had studied as a gradu- ate student or even the variety of species I had col- lected in the Amazon region of South America. In recent years (including the time I am writing this), my house in Ibadan, Nigeria, allows me to go out every evening and watch spectacular, huge clouds of flying foxes migrating overhead from their daytime camp in an urban, botanical garden, where they are safe from hungry hunters, to their evening fruit-tree feeding grounds. The flying foxes are fruit and nectar eating bats of the suborder Megachiroptera, which are found in the old world tropics and are only distantly related to the suborder Microchiroptera which I studied as a stu- dent. This book did confuse me a bit, at first, since the authors primarily discuss only Australian species and the most common of these in the genera Pteropus, which does not occur in continental Africa. The authors state in the introduction “There is a number of other fruit-eating Megachiroptera which are not in the genus Pteropus but which are also called ‘flying foxes’. ” Then throughout the rest of the book they talk of flying foxes as if they were only Australian Pteropus. This caused me consider- able confusion since I have been involved in some flying fox research in Nigeria, and I know this is not Australia. Back to my early lessons about how Latin, scientific names are so much more precise than com- mon names. This book is very comprehensive, well re- searched, and well written. It provides excellent information to those who are interested in flying foxes, even if not just in Australia. It covers such areas as history and archaeology, identification, and distribution of the Australian species, anatomy and physiology, disease, behaviour, feeding ecolo- gy, migration, conservation, rehabilitation, and even hand rearing, in great detail. Appendices include native plant foods, extensive references, a glossary, and index. There is some confusion where references in the text to an individual or institution doing research cannot be traced to the specific published references at the back of the book. Some very interesting facts are presented in this book. Did you know that bats are the second most common mammal group to rodents. Then there is the blossom bat, described as a “mouse-sized flying fox” — a bit of a contradiction. Flying foxes, unlike other bats, rely largely on sight. They will not fly in total darkness and do not hear much better than a human being. Their vision is similar to that of a cat. During the day, flying foxes roost together in large “camps” which can number over 250 000. These camps can be very noisy and result in damages to the local trees. At night, they will migrate long distances to feed, again causing damages due to their numbers. This has resulted in cropping by orchard owners, which has decimated the population in Australia. They are now protected and conserved for their ben- eficial behaviours such as pollination and dispersal of fruit seeds. Flying foxes have learned the behaviour of roost- ing in urban forests or parks as a protection against shooting. In my area of Nigeria, the largest local camps are the botanical garden close to our house, and two local university campuses. Flying foxes are considered a culinary delicacy in Nigeria as well as elsewhere in their range. One of my co-workers here says they make delicious soup. But he was surprised that they do not taste like a bird. Usually if I eat bush meat, I do not ask what it is. With a lifespan (in captivity) of up to 50 years, flying foxes only produce one young per year. This has resulted in serious population drops due to habitat loss and hunting pressure. Several southeast Asian islands have seen populations become extinct 168 THE CANADIAN FIELD-NATURALIST and the entire population of some species in Australia is now smaller than the 300 000 reported in one camp less than fifty years ago. Another species which I hope will not disappear before we have really had the chance to get to know it. This book is a very worthwhile and authoritative source BOTANY Vol. 116 of information on the intriguing world of flying foxes. WILSON EEDY Geomatics Nigeria Limited, Plot 7 Off DPC Road, Ibadan, Nigeria Guide to Standard Floras of the World: An Annotated Geographically Arranged Systematic Bibliography of the Principal Floras, Enumerations, Checklists, and Chorological Atlases of Different Areas. Second Edition. By David G. Fordin. 2001. Cambridge University Press, The Edinburgh Building, Cambridge CB2 2RU UK; 40 West 20th Street, New York, NY 10011-4211, U.S.A. xxiv + 1100 pages U.S. $240.00 + $6.00 postage. This volume, which includes information on all the floras of the world known to the author up to the end of the twentieth century, is a tremendous step ahead of the first edition which was published in 1984 and numbered 619 pages. The first pages of this volume include prologues and acknowledge- ments of the first and second editions; Part I which numbers 84 pages and includes an analytical- synthetic systematic bibliography of “standard” flo- ras: scope, sources and structure, the evolution of floras, floras at the end of the twentieth century: phi- losophy, progress, and prospects and references. Part II includes systematic bibliography, conspectus of divisions and superregions, world floras, isolated oceanic islands and polar regions, and divisions: North America (north of Mexico), Middle America, South America, Australia and islands of the south- west Indian Ocean (Malagassia), Africa, Europe, northern, central and southwestern (extra-monsoon- al) Asia, southern, eastern and southeastern (mon- soonal) Asia, Greater Malasia, and Oceania. This is followed by Appendix A: major general bibliogra- phies, indices, and library catalogues covering world floristic literature, and Appendix B: Abbreviations of serials cited, addenda in proof, geographical index, and author index. The main part of this volume then proceeds through the areas listed in Part II above with descriptions of the regions, locations, number of species, early botani- cal history up to the end of the twentieth century, and the major floras which have been published through the years. There is a wealth of information here both for those living in any region or anyone interested in the botany of another part of the world. It is inevitable in a work such as this that occa- sionally some useful publications will be passed by. Some from northern North America that might have been included are the following: Benson, L. 1982. The Cacti of the United States and Canada. Stanford University Press. Biek, D. 1999. Flora of Mount Rainier National Park. Oregon State University Press. Cody, W. J. 1988. Plants of Riding Mountain National Park, Manitoba. Agriculture Canada. Douglas, G. W., C. B. Straley, and D. V. Meidinger. 1998. Rare Native Vascular Plants of British Columbia. Crown Publications. Hallworth, B., and C. C. Chinnappa. 1997. Plants of Kananaskis Country in the Rocky Mountains of Alberta. University of Calgary Press. Holmgren, A. H. 1998. “Illustrated Companion to Gleason and Cronquists’ Manual: Illustrations of the Vascular Plants of Northeastern United States and Adjacent Canada”. The New York Botanical Garden. McGregor, R. L., and T. M. Barkley. 1977. Atlas of the Flora of the Great Plains. The Iowa State University Press. Packer, J., and C. E. Bradley. 1984. Checklist of the rare plants in Alberta. Provincial Museum of Alberta National History Occasional Paper Number 5. Petrik-Ott, A. J. 1979. The Pteridophytes of Kansas, Nebraska, South Dakota and North Dakota. Nova Hedwigia, Beiheft 61. Argus, George [Series Editor] 1977-1995. Rare Native Vascular Plants of Canadian Provinces and Territories. Syllogeus National Museum of Natural Sciences, Ottawa. Tryon, A. F., and R. C. Moran. 1997. The Ferns and Allied Plants of New England. Centre for Biological Conservation, Massachusetts Audubon Society. Botanical studies will continue around the world and already one for the year 2000 has been published in Canada: H.R. Hinds, Flora of New Brunswick, Second Edition, which was a great step ahead from the First Edition. It will be most important to keep The Guide to Standard Floras of the World up to date through new editions or supplements. WILLIAM J. Copy Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada. 2002 BOOK REVIEWS 169 EVIRONMENT Their Fathers’ Work: Casting Nets with the World’s Fishermen By W. McCloskey. 2000. International Marine/McGraw- Hill, New York. 370 pp. U.S. $20.95 This book provides the reader with a superb and highly praised overview of global fisheries, focusing on Alaskan waters. In addition, it also covers first- hand experiences for offshore and coastal fisheries with vessels from Japan, Chile, Indonesia, New- foundland (Grand Banks), Maine (Georges Bank), Iceland, and Norway. The book is very pleasant to read as it combines fiction with fact. It is a heroic and romantic description of a likely soon-to-be-gone life of hard work. Nevertheless, reading how other people work very hard and under life-threatening conditions might present some sort of decadence; but so be it. In case the reader is not familiar with how to cheat in this business of international fisheries and quotas (led by Spain, Taiwan, Japan, and many East European nations) this book will be enlightening. It outlines in detail how fishing quotas are easily dou- bled, if not ignored, by many vessel captains and fishermen worldwide. The explicit use of Dynamite Fishing, Liner Nets (an additional net with an ille- gally smaller mesh-size put inside the regular net), the “Pareja” Method (one huge net pulled by two boats), and many other tricks are shown and suggest- ed: e.g., the same vessel being registered with two different names (thus, multiplying the quota by two), stowing an additional catch somewhere under deck, trading the catch offshore (therefore enabled to start again with a “new” quota), and mis-reporting catch- es. When fisheries officers appear for control and gear inspection, nets simply get cut off (which makes it even worse for fish, seabirds, and sea mam- mals that drown in the “ghost nets” later). Overall, I find that the author, an American, might have a ten- dency to put too much blame on the Spaniards and Russians. Instead, a mention and description of the role that the Vladivostok-based Russian fisheries play, acting worldwide, could have made the book even better. The thorough understatement of environmental damage done by coastal and offshore fishery must be of concern to any informed naturalist. The author neglects to address the destructive fishery method from draggers (“seafloor dredging’’), which is esti- mated to damage an area larger than that lost through deforestation in the tropics. There is no mention of fisheries gear polluting beaches world- wide, or “ghost nets” which float around the world’s oceans for years (eventually, they will sink, but only the fish know whether they will ever rot). Sensitive by-catch topics such as the endangered Short-tailed Albatross (Phoebastria albatrus) caught by freezer-longliners fishing off Alaska are not mentioned, and certainly there is no reporting of the numerous sea turtles, sharks, dolphins, porpois- es, seabirds, moon fishes, and many other species suffering and dying for the sake of high quality fish. In times of environmentalism, that might be seen as a short-coming of this book. Although the occur- rence of a “black catch” is somewhat mentioned, one reads that shrimp fisheries have apparently almost no by-catch. The reader has to hold his/her breath when McCloskey mentions “overpopula- tions” of Sockeye and seals; 50 000 seals are described as an “overpopulation” rather than vic- tims in a potential by-catch problem. No wonder, the author identifies clearly from the “fishermen’s side”, blames Greenpeace, and does not place fish- eries in the overall context of the environment. Instead, he largely focuses on economical and descriptive aspects of fisheries. In this regard, the author’s presentation of Chile’s fishery develop- ment lacks sensitivity to the well-proven and nega- tive effects of over-commercialization. On the other side, his wonderful and detailed presentation of the effects from the Exxon Valdez Oilspill for Alaska and its island communities compensate for the pre- vious short-comings. A remarkable link is shown why the prizes of the Japanese salmon market are driven by cycles of the Japanese salmon runs and thus dictate the Alaskan salmon fisheries. McCloskey gets closer to the heart of the fisheries problem when outlining that improved efficiency and introduction of very light, and therefore allow- ing for longer, plastic nets has contributed to the current overfishing crisis. In the numerous and fascinating chapters the author also emphasizes and describes that there exists such a thing as severe overfishing: Snow Crab in Alaska; Cod, Flounder and Squid in Newfoundland; and Halibut off West America. He blames governmental mis-management and elaborates nicely throughout the text that there is also conflict of interest among fisher- men on these topics; e.g., unions, and small scale fish- eries vs. industrialized trawlers. In the context of gov- ernmental mismanagement, New Zealand’s Orange Roughy, a prime example of overfishing and disas- trous fisheries management, could have been men- tioned, too. The book would have been even better if topics such as a Native Fishery Rights, North Sea Fisheries, and Krill Fisheries in the Antarctic had been included. The map of the Grand Banks lacks the French Fisheries zone around St. Pierre and Miquelon; but the reader will appreciate that this book has a very detailed index, which allows that it can serve as a valid source of references. The book ends with a well-written and conclu- sive section on global fisheries and policy. The author quotes from one of his many interviews with 170 THE CANADIAN FIELD-NATURALIST experts: “Gathering fishery statistics is an art in probability”. That statement makes it clear that, cur- rently, there can be no sustainable world fisheries. Due to the many topics covered, I thoroughly enjoyed reading this book and got literally “hooked”. AAAS Atlas of Population and Environment By P. Harrison and F. Pearce. 2001. University of California Press, Berkeley. xi + 204 pp., illus. Cloth U.S. $65; paper U.S. $29.95. Population is the main cause of environmental problems. This book provides an excellent, well- documented, and authoritative demonstration of this principle, using easy to read text and great atlas dis- plays. It needs to be read by every person who is concerned over our world environmental degradation and the population impacts of the very near future. It comes home to me personally, spending much time in Nigeria, one of the highest and most densely pop- ulated countries in Africa. In spite of the HIV-AIDS epidemic, it still has one of the highest population growth rates in the world (2.81% per year). Here one comes face to face with poverty, lack of clean water supplies, major losses of forest and ecological habi- tat, desertification, and the desperation of many peo- ple struggling to survive from day to day. I see things that I never would have believed before I first came here 10 years ago. What will it be like in 2050 AD when the population is expected to grow by more than 50% to about 200 million people (in an area the size of Ontario, with large areas of arid savanna). This book gives some predictive outlook for such areas of the world, where the major popula- tion growth will be in the future. I do not think this is an issue that we can afford to ignore. Perhaps, as a bit of a field-naturalist heretic, I suggest that it is more of an environmental priority than the birds in our back yard. I recommend this book as a critical reference, resource for anyone concerned about the environment and the future of our world. The book is organized into three major sections. Part 1: Overview (40 pages) is primarily introducto- ry text, covering issues such as environmental poli- cies and philosophies, the scale of our existence, and the theory of population-environment links. How many people can the world sustain? This area is not well understood and the answer is not given, but the- ories such as those of Malthus and Erhlich are ana- lyzed in an impartial scientific manner. Much data is presented, but the reader is often left to make their Vol. 116 FALK HUETTMANN Centre for Wildlife Ecology, Biology Department, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia V5A 1S6 Canada own conclusions. In the last century population expansion has occurred at an alarming, unprecedent- ed rate (from three million — 2000 years ago, to 1 billion 200 years ago, to over 6 billion now, and pre- dictions to over 9 billion by 2050). In the last centu- ry, we have lost/of the world’s topsoil, 1/5 of our agricultural land base, major parts of our forest, and other habitat. We have experienced a species extinc- tion rate 100 times that of any time in our previous history. Our problems are summarized in a rather sickening and scary manner, but easily understood by those of us who live much of our lives in the third world. The second part of the book is 164 pages of an atlas of resources and resource uses or losses, along with some accompanying discussion and references. Areas covered include population effects on natural resources, land use, atmosphere, wastes and chemi- cals, ecosystems, and biodiversity. The atlas maps allow easy reference and the text provides excellent detail. The third part of the book provides a few case his- tories of population-environment interaction, for dif- ferent biogeographic regions of the world. All but one are taken from World Wildlife Fund informa- tion. The one exception, and location closest to Canada, is a Nature Conservancy discussion of the Sonoran Desert (a favourite retreat for me). I guess Canada can be happy to be on the good side of most of these environmental issues and to be proud not to have a case history illustrated here. However, I do strongly believe that we cannot hide from these potential future population-caused envi- ronmental catastrophes. We are all in one world and the information in this book is an excellent start to planning to save it. WILSON EEDY Terfa Inc, RR# 1, Glencairn, Ontario LOM 1K0O Canada and Geomatics Nigeria Limited, Plot 7 Off DPC Road, Ibadan, Oyo State, Nigeria (terfa@geoniger.com, www.geoniger.com) 2002 BOOK REVIEWS 17t Making Better Environmental Decisions By Mary O’Brien. 2000. The MIT Press, Cambridge, Massachusetts. 286 pp., illus. U.S. $22.95. This book starts with a story, a parable of sorts. A woman is standing by an icy mountain river, intend- ing to cross to the other side. A team of risk asses- sors accompanies her, reviewing her situation. They present arguments and calculations, concluding that the risk she faces is low from their various perspec- tives — toxicology, cardiology, hydrology, policy — and that she should wade across the river. Yet, to their extreme frustration, she refuses to wade. Why? She points upstream and tells them, “Because there is a bridge.” The risk assessors evaluated the risk of only one option: wading across the river. The woman reviewed her alternatives, which included crossing on the bridge upstream. Making Better Environ- mental Decisions is about doing what the woman in the parable did: making decisions based on all possi- ble alternatives instead of considering only the risk associated with a narrow range of options. The concept of alternatives assessment is based on certain common-sense principles. These include, among others, that it is unacceptable to harm human Scientific Method for Ecological Research By E. David Ford. 2000. Cambridge University Press, Cambridge. 564 pp., illus. Cloth U.S. $10; paper U.S. $49.95 Dr. Ford, professor, College of Forest Resources, University of Washington, carves out, in exemplary fashion, a philosophy of ecological research. The philosophical exploration of the “hard” physical sci- ences as the paradigmatic sciences introduced assumptions as to what makes “good science,” but it has become evident that certain of these assumptions are questionable or even false when it comes to the biological sciences and ecology. The philosophy of the physical sciences has created a “storybook scien- tist’ who is considered specialized, reductionist, sci- entifically objective, rigorous, experimental, predic- tive, and essentially value-free. For Professor Ford, such a description is a caricature. Ford eschews any scientific method which defies particular techniques, a particular form of scientific reasoning, asserts that cer- tain questions are more worthwhile than others, or assumes that questions must be asked in specific ways. For the author, the scientific method is not an immutable, monolithic process characterized by true and tried roadsides, but a creative process in which these roadsigns may take many and varied forms depending on the particular research question at hand. Written with the graduate student and aspiring sci- entist in mind, Ford’s weighty tome is broken down or non-human life when reasonable alternatives exist. That nobody should be able to define for any- one else how much damage is “acceptable”. That those affected by decisions should be involved in making them. And that there are always alternatives that can be considered. The book outlines why risk assessment — the pro- cess currently relied on by governments and the pri- vate sector to make decisions — doesn’t work, and goes on to explain why alternatives assessment is bet- ter and how to make the shift. Clearly written and easy to read, the book is full of convincing examples, including many relating to ecosystems and wildlife, that speak loudly for considering all possible alterna- tives before making decisions that will affect the health of humans and the natural environment. This book is a must-read for all naturalists involved in decisions-making processes, as profes- sionals or as volunteers, and for anyone concerned about the future of life on this planet. R. SANDER-REGIER RRS5 Shawville, Quebec JOX 2Y0 Canada into four main sections; an analytical framework for conducting research, the basis for scientific inference and theory development in ecology, the social dimensions of research, and finally the presentation of a methodology for developing ecological theory, his so-called progressive synthesis. Making explicit how ecologists think in their research is his over- arching goal. Of significance to Ford is a recognition of the unique features of ecological research which demand a research methodology different from that assumed for the physical sciences. Of particular con- cern is the open or partially open nature of ecologi- cal systems, the probabilistic nature of many ecolog- ical concepts, the inherent variability of ecological data, and problems of multiple causality. All of these factors pose greater difficulties in measurement and concept development, conditions for scientific infer- ence, and the basis for theory acceptance or refu- tation when compared to controlled, laboratory conditions common to the physical sciences. Ford’s pedagogically refined review of the diverse forms of scientific reasoning and philosoph- ical assumptions grounding scientific research is exhaustive and brimming with insight. Lack of space prevents the consideration of many critical issues, but attention to a select few is profitable. Particular emphasis is placed by Ford on the critical analysis of concepts and propositions, both of which Ee 172 THE CANADIAN FIELD-NATURALIST will act as the foundation of research. What is known (axioms), what one wishes to know (postu- lates), and how one intends to collect data that will assess postulates (data statements) are critical first steps in research. The clarification of ecological concepts is important as well. Ecology often focuses on what Ford terms integrative concepts (e.g., sta- bility, ecological integrity, resilience), theoretical constructs about ecological organization that defy direct measurement and demand synthesis from a variety of system studies. Caution is directed to the uncritical use of statistical inference as the sole assessment of postulates. Highly significant is the notion of theory domains which define the bound- aries within which a particular theory may or may not operate, effectively rendering ecological theory development very complex. Of significance as well is the notion that both measurement and experiment are essentially an art. Even when precise and accu- rate, all measurements by definition are abstracts in that they represent the object of interest, but are not the things themselves. Ford refutes the notion that Popperian falsification can be used as the basis for theory change, citing evidence to show that the rational basis for theory acceptance or rejection 1s much more complex. The reader is introduced to the “subjective” or sociological dimensions of science. Science is pre- sented as not simply an automated, invariant or determined process, but as a human activity under- MISCELLANEOUS The Last Great Sea: Vol. 116 taken by men and women acting as bearers of wis- dom, passion, faults and foibles seemingly no more or no less than the “average citizen.” Ford addresses such issues as scientific fraud, the publication pro- cess, peer-review, the significance of gender, and the role of scientists in developing public policy. Social processes are viewed as significant in determining which questions or research problems are considered important, which group will receive funding resources and precisely how research questions are formulated and pursued. A popular view of science may assume that the scientific enterprise is a wholly “objective” process to which the particular investigator simply submits. Professor Ford returns a human face to science emphasizing the role of creativity, choice, probabili- ties, the art of science, plurality of logical reasoning and methodological ideals, a distancing from a priori philosophical assumptions. This by no means weak- ens the scientific enterprise, but rather re-introduces a rigorous and critical attitude in posing scientific problems and developing an adequate procedure of assessment. In doing this, Professor Ford ennobles human scientific activity by calling explicit attention to the pattern of scientific reasoning. JOHN McCarthy, S. J. St. Mark’s College, University of British Columbia, 5935 Iona Drive, Vancouver, British Columbia V6T 1J7 Canada A Voyage through the Human and Natural History of the North Pacific Ocean By T. Glavin. 2000. David Suzuki Foundation and Greystone Books, Douglas & McIntyre Publishing Group, Vancouver/Toronto. 244 pp. This book belongs in every conservationist’s bookshelf, to say the least. As D. Suzuki describes very convincingly in the foreword, the environment of today’s North Pacific is characterized by its loss of (fish) species and its wipe-out of protein assem- blages. The collapse of Sockeye Salmon is only one of many sad examples, many more exist: Steller’s Sea Cow, Spectacled Cormorant, Dwason’s Caribou (Queen Charlotte Islands), and even plant species like Tobacco (Queen Charlotte Islands). Other species like Walruses, Sea Otters, and Fur Seals have barely survived until now. The first chapter starts slowly but allows a very solid overview about historical and archaeological facts. After Chapter 2, nobody can deny anymore the environmental disaster and mis-management of the North Pacific and coastal British Columbia. Nevertheless, the author convinces the reader that the North Pacific still is THE largest fish producer in the world. “As in aboriginal fisheries, mythology played a part in industrial fisheries management, especially the myth of a superabundant ocean and the all-powerful capability of science and technology to fix the messes made by hydroelectric dams, lousy forestry practices and overfishing”. The governmentally encouraged Merganser Control and Bear Shooting Programs designed for the sake of Salmon Protection prove this citation very well. Galvin strongly eliminates all illu- sions on how to heal the problem of overfishing. For instance, he shows that S. Livingstone’s widely fol- lowed idea of Fish Hatcheries does not produce more salmon, but instead takes away funds and harms natu- ral salmon stocks since they simply replace the last remaining and struggling stocks with poorly adjusted new ones; Gavin’s arguments are also strong against 2002 BOOK REVIEWS 173 Salmon Farming; e.g., it contributes to the closure of marine fisheries for wild salmon, and it requires 3 kg of fish to produce 1 kg of salmon. All Pacific salmon species are discussed: Chum, Sockeye, Pink, Coho, Steelhead, Masu and Amago. Of major interest is in this regard the scientific dis- cussion around the taxonomy of salmon; e.g., Steelhead (classified until 1980s as Trout). The author brilliantly points out the implications of the religious-based and somewhat outdated taxonomical system by Carl von Linné, and how this affects the species management by national governments (provincial and federal) on an international level even (Canada vs. U.S.A). The backwardness and failure of fishery laws are shown by outlining that the first salmon-fishing reg- ulations for the Fraser River was a simple word-for- word replication of fishing regulations on English Rivers. At that time, Canada’s external affairs juris- diction was still controlled by the British, which affects the Canada-U.S. salmon treaty concluded 1930s and renewed in 1985. In addition, Galvin shows that Canadian and U.S. fishery scientists sig- nificantly differed in their stock assessment results for the same species in the same waters even; conse- quently, so did the management and political agen- das. This is the classical picture of “mixed-stock” fisheries, which also threatens small salmon runs. The author reports the incidental death of 50 000 marine mammals and 500 000 seabirds due to driftnet fishery activities in the North Pacific; marine (plastic) pollution comes with it. Despite the well shown fail- ure of a European and Western approach dealing with the North Pacific fisheries, domestic Japanese, and Native fisheries seemed to work well and sustainable. Galvin shows the magnitude of “pre-contact” fishery for salmon by natives, which was even comparable with levels of commercial fisheries from this century. Some readers might find that the book slightly fol- lows stereotypical views of the noble native. A very strong point in this book is how the North Pacific and its fauna is linked with the “hinterland”: Old-growth rainforest, landscape and Bald Eagles. This needs to be considered in the light that resident Killerwhales in British Columbia are among the most contaminated cetaceans of the world. A very complete picture of the North Pacific is portrayed by fully considering the Russian influence and history. The book outlines well that Russian set- tlers did much better than the western type of colo- nization (a point that might be put in doubt for the Kodiak Islands at least). The Russian-American Company was much more relevant in the history of North Pacific settlements and explorations than the Hudson Bay Company (HBC). But nevertheless, as with the HBC, the Russian quest for the North Pacific had the same motivation: central European pelt resources were already overhunted! Regarding the marine ecology of the North Pacific, the importance of the Aleutian low, Pacific currents, and El Nino are fully described. This ecosystem is driven by ‘regime changes’, which calls for a dynam- ic management. The author outlines this very well by presenting the ground-breaking work from Russian Scientist T. Baranov, but also from Bill Ricker (“Ricker curve’’) and others at the Pacific Biological Station; e.g., G. McFarlane and D. Beamish. A quote from the book says it all: Understanding catch statis- tics is like “reading a single faded and crumbling onionskin page from an early draft of Wagner’s Tannhaeuser, in a dimly lit room”. Another quote of the book and taken from the U.N. Code of Conduct for Responsible Fisheries states, in part, that “the absence of adequate scientific information should not be used as a reason for postponing or failing to take conservation and management measures’. Well said. The chapters on anthropology and human history of the North Pacific and how the Russians, Asians, and Natives settled and explored the North Pacific are on the same level than high-caliber books as Guns, Germs and Steel by J. Diamond. Just to name some highlights, Glavin mentions how natives grew Arrowhead and potatoes, he cites the work of the Russian Anthropologist S. Fedorova, and he docu- ments that Hawaiians, Japanese, Chinese and Russians presented a major group of settlers. In addi- tion, the book reports a lot of British Columbian and Vancouver history and puts Canada in the context of the overall Pacific. Despite the fact that whaling, sealing, and eating dolphins are as old as the human history of the North Pacific, whale watching (starting as early as 1907) has already produced more profit than commercial whaling ever did for western North America. Green- peace started in Vancouver; it “was born in the blood of whales”. Nevertheless, the author shows that already in the 19th century the pelagic seal hunt pro- voked the first great international controversy about the overharvesting of the world’s marine mammals. It resulted in the international milestone contract (‘fur seal treaty’) of 1911 between Russia, Japan, Canada and U.S.A. FALK HUETTMANN Centre for Wildlife Ecology, Biology Department, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia V5A 1S6 Canada 174 THE CANADIAN FIELD-NATURALIST NEw TITLES Zoology Animal ecology. 2001. By C. Elton. University of Chicago Press, Chicago. xvii + 209 pp., illus. U.S.$20. *A birder’s guide to metropolitan areas of North Amer- ica. 2002. Edited by P. Lehman. American Birding Association, Colorado Springs. iv + 508 pp., illus. +The birds of heaven: travels with cranes. 2001. By P. Matthiessen. Greystone Books, Douglas and McIntyre, Vancouver. xv + 349 pp., illus. $36.95. *The complete guide to the birds of Europe. 2000. By K. Mullarney, L. Svsnsson, D. Zetterstrom, and P. J. Grant. Princeton University Press, Princeton. 399 pp., illus. U.S.$49.50. +Conversations with an eagle: the story of a remarkable relationship. 2002. By B. Cox. Greystone Books (Douglas and McIntyre), Vancouver. 288 pp. $22.95. *Coral reef fishes. 2002. By E. Lieske and R. Myers. Revised edition. Princeton University Press, Princeton. 400 pp., illus. U.S.$24.95. *The Cuban treefrog in Florida. 2001. By W. E. Meshaka, Jr. University Press of Florida (Canadian distributor Scholarly Book Services, Toronto). xxiii + 191 pp., illus. *Dinosaurs: the encyclopedia, supplement 2. 2002. By D. F. Glut. McFarland, Jefferson, North Carolina. x + 685 pp., illus. U.S.$75. The family butterfly book: projects, activities, and a field guide to 40 favorite North American species. 2001. By R. Mikula. Storey Books, Pownal, Vermont. x + 166 pp., illus. Cloth U.S.$29.95; paper U.S.$16.95. +Geographic variation in size and shape of savannah sparrows (Passerculus sandwichensis). 2001. By J.D. Rising. Cooper Ornithological Society, Camarillo, Cali- fornia. 65 pp., illus. U.S.$7. The grizzly almanac. 2001. By R. H. Busch. Lyon Press, New York. 240 pp., illus. U.S.$29.95. *Guia de las aves de Espana. 2000. By E. de Juana and J. M. Varela. Lynx Edicions, Barcelona. 223 pp., illus. 15 Euros. *Handbook of the birds of the world: volume 7: jaca- mars to woodpeckers. 2002. Edited by J. de Hoyo, A. Elliot, and J. Sargatal. Lynx Edicions, Barcelona. 613 pp., illus. U.S.$185. *National Audubon Society guide to marine mammals of the world. 2002. By R. R. Reeves, B. S. Stewart, P. J. Clapham, and J. A. Powell. Alfred A. Knopf, New York. 527 pp., illus. U.S. $26.95. Canadian distributor Random House, Toronto. $39.95. Neotropical treeboas: natural history of the Corallu hor- tulanus complex. 2002. By R. W. Henderson. Kreiger Publishing, Melbourne, Florida. 228 pp., illus. U.S.$44.50. Vol. 116 New animal discoveries. 2001. By R. Orenstein. Twenty- First Century Books, New York. 64 pp., illus. U.S.$25. Noises in the night: the habits of bats. 2001. By D. Kovacs. Steck-Vaughn, Austin, Texas. 48 pp., illus. US: $25: Pheasants, partridges, and grouse. 2002. By S. Madge and P. McGowan. Princeton University Press, Princeton. 480 pp., illus. U.S. $49.50. A pocket guide to reptiles and amphibians of Alberta. 2002. By A. P. Russell and A. M. Bauer. Red Deer Press, Calgary. 260 pp., illus. $24.95. Sea soup: zooplankton. 2001. By M. M. Cerullo. Til- bury House, Gardiner, Maine. 40 pp., illus. U.S. $16.95. *Sharks. 2002. By A. and A. Ferrari. Firefly Books, Willowdale, Ontario. 256 pp., illus. $24.95. Tadpoles of south-eastern Australia: a guide with keys. 2002. By M. Austis. Kreiger Publishing, Melbourne, Florida. 281 pp., U.S.$44.50. *Tales from the underground: a natural history of sub- terranean life. 2001. By D. W. Wolfe. Perseus, Cam- bridge, Massachusetts. x + 223 pp., illus. U.S.$26. +Thunder on the tundra: Inuit qaujimajatugangit of the Bathurst caribou. 2002. By N. Thorpe, N. Hakongak, S. Eyegetok, and the Kitikmeot Elders. Available through Natasha Thorpe, 231 Irving Road, Victoria, B.C. V8S 4A1. 208 pp., illus. $42. +Turtles and tortoises. 2002. By V. Ferri. Firefly Books, Willowdale, Ontario. 256 pp., illus. $24.95. Why elephants have big ears: understanding patterns of life on earth. 2001. By C. Lavers. St. Martins, New York. 288 pp., illus. U.S.$23.95. Botany *Dictionary of the fungi. 2001. By P. M. Kirk, P. F. Cannon, J. C. David, and J. A. Staples. CAB International (distributed by Oxford University Press, Cary, North Carolina). xi + 655 pp., illus. A field guide to water and wetland plants of the prairies. 2002. By H. Lahring. Red Deer Press, Calgary. 192 pp., illus. $18.95. {+Forest dynamics and disturbance regimes: studies from temperate evergreen — deciduous forests. 2002. By L. E. Frelich. Cambridge University Press, New York. ix + 266 pp., illus. U.S.$70. Harmful algal blooms on the North American west coast. 2001. Edited by R. Ralonde. Alaska Sea Grant College Program, Fairbanks. 73 pp. U.S$10. 2002 +New world botany. 2001. By R.H. Peterson. Koeltz Scientific Books, Koenigstein, Germany. xv+ 638 pp., illus. Cloth Euro 93; paper Euro 62. Wildflowers around the year. 2001. By H. Ryden. Clarion, New York. 90 pp., illus. U.S.$17. Wild flowers of field and slope. 2002. By L. Clark. Harbour Publishing, Madeira Park, British Columbia. 80 pp., illus. $9.95. Environment The B.C. roadside naturalist. 2002. By R. Cannings and S. Cannings. Greystone Books (Douglas and McIntyre, Vancouver). 240 pp., illus. $29.95. The beachcombers’s guide to seashore life of Cali- fornia. 2002. By J. D. Sept. Harbour Publishing, Madeira Park, British Columbia. 300 pp., illus. CAN $28.95; U.S.$17.95 +Conserving living natural resources in the context of a changing world. 2002. By B. J. Weddell. Cambridge University Press, New York. xvi + 426 pp., illus. U.S.$35. Evolution’s workshop: God and science on the Galapagos Islands. 2001. By E. J. Larson. Basic Books, New York. 322 pp., illus. U.S.$27.50. The functional consequences of biodiversity: empirical progress and theoretical extensions. 2002. Edited by A. P. Kinzig, S. W. Pacola, and D. Tilman. Princeton University Press, Princeton. 392 pp., illus. Cloth U.S.$75: paper U.S.$29.95. Green phoenix: restoring the tropical forest of Guanacaste, Costa Rica. 2001. By W. Allen. Oxford, New York. 310 pp. U.S.$35 7Introduction to conservation genetics. 2002. By R. Frankham, J. D. Ballou, and D. A. Briscoe. Cambridge University Press, New York. xi+ 617 pp., illus. Cloth U.S. $130; paper U.S.$50. +Life at the limits: organisms in extreme environments. 2002. By D. A. Wharton. Cambridge University Press, New York. xi + 307 pp., illus. U.S.$25. *Making parks work: strategies for preserving tropical nature. 2002. Edited by J. Terborgh, C. van Schaick, L. Davenport, and M. Rao. Island Press, Washington. xix + 511 pp., illus. Cloth U.S.$65; paper U.S.$32.50. Marine community ecology. 2001. Edited by M. D. Bertness, S. D. Gaines, and M. E. Hay. Sinauer, Sunder- land, Massachusetts. 550 pp., illus. U.S.$59.95. *Multitrophic level interactions. 2002. Edited by T. Tschrntke and B. A. Hawkins. Cambridge University Press, New York. vii+274 pp., illus. U.S.$75. *Politics of the wild: Canada and endangered species. 2001. Edited by K. Beazley and R. Boardman. Oxford BOOK REVIEWS 175 University Press Canada, Toronto. x + 254 pp., illus. $27.95. 7Reef life. 2002. By A. and A. Ferrari. Firefly Books, Willowdale, Ontario. 288 pp., illus. $24.95. 7+Spread sheet exercises in conservation biology and landscape ecology. 2002. By T. M. Donovan and C. W. Welden. Sinauer Associates, Sunderland, Massachusetts. xi + 464 pp., illus. U.S.$5. 7Spread sheet exercises in ecology and evolution. 2002. By T. M. Donovan and C. W. Welden. Sinaeur Associates, Sunderland, Massachusetts. xi + 556 pp., illus. U.S.$5. y+Summary of knowledge acquired in northern environ- ments from 1970 to 2000. 2001. By G. Hayem. Hydro- Quebec, Montreal. 110 pp., illus. Free. *Two hundred years of ecosystem and food web changes in the Quoddy Region, Outer Bay of Fundy. 2002. By H. Lotze and I. Milewski. Conservation Council of New Brunswick, Fredericton. The tropical rainforest: explore the natural world of the rainforest swamplands and the interior. 2001. By G. Cheshire. Crabtree, New York. 39 pp., illus. U.S.$18. Visions of the wild: a voyage by kayak around Van- couver Island. 2001. By M. Coffey and D. Goering. Harbour Publishing, Madeira Park, British Columbia. 182 pp., illus. $36.95. +Weird nature: an astonishing exploration of nature’s strangest behavior. 2002. By J. Downer. Firefly Books, Willowdale. 156 pp., illus. Cloth $35; paper $19.95. Wild solutions: how biodiversity is money in the bank. 2001. By A. Beattie and P. Ehrlich. 240 pp. U.S.$25.95. Miscellaneous +Catalogue of meetings 1988 to 2001 MclIlwraith Field Naturalists. 2002. By W. W. Judd. Phelps Publishing, London, Ontario. 28 pp. $8. *Cheltenham in Antarctica. 2002. By D. M. Wilson and D. B. Elder. Reardon Publishing, Cheltenham, England. 140 pp., illus. E9.99. *+Common and contested ground: a human and environ- mental history of the northwest plains. 2002. By T. Binnema. University of Oklahoma Press, Norman. xvi + 263 pp., illus. U.S.$29.95. *The one culture: a conversation about science. 2001. Edited by J. A. Labinger and H. Collins. University of Chicago Press, Chicago. xii+ 329 pp. U.S.$18. *On her own terms: Annie Montague Alexander and the rise of science in the American West. 2001. By B. R. Stein. U. California P., Berkeley. xvii +380 pp., illus. U.S.$35. 176 Books for Young Naturalists About amphibians: a guide for children. 2001. By C. Sill. Peachtree, Atlanta. 40 pp., illus. U.S.$14.95. Animals nobody loves. 2001. By S. Simon. North-South Books, New York. 48 pp., illus. U.S.$15.95. Bears. 2001. By D. Ferte, M. Reddy, and E. D. Stoops. Sterling, New York. 80 pp., illus. U.S.$17.95. Caimans. 2001. By S. Dollar. Raintree Steck-Vaughn, Austin. 32 pp., illus. U.S.$23. Dolphins. 2001. By J. Vogel. NorthWord Press, Minnetonka, Minnesota. 48 pp., illus. U.S.$7.98. Global Warming: the threat of earth’s changing cli- mate. 2001. By L. Pringle. North-South Books, New York. 48 pp., illus. U.S.$16.95. Kids care for the earth. 2002. By G. Thompson. National Geographic Society, Washington. 32 pp., illus. U.S.$41.95 (pack of 6). Manatees. 2001. By K. Feeney. NorthWord Press, Minnetonka, Minnesota. 48 pp., illus. U.S.$7.95. 100 award — winning science fair projects. 2001. By G. Vecchione. Sterling, New York. 208 pp., illus. U.S.$21.95. THE CANADIAN FIELD-N ATURALIST Vol. 116 Protecting our planet : animal watch. 2001. By R. Few. DK Publishing, New York. 60 pp., illus. U.S.$16.95. Rain forest. 2001. By E. Greenwood. DK Publishing, New York. 48 pp., illus. U.S.$9.95. Recycling and reuse: our impact on the planet. 2002. By R. Bowden. Raintree Steck-Vaughn, Austin, Texas. 64 pp., illus. U.S.$18.98. The seashore. 2001. By A. Wilkes. Kingfisher, New York. 32 pp., illus. U.S.$11.95. Sharks. 2001. By L. Evert. NorthWord Press, Min- netonka, Minnesota. 48 pp., illus. U.S.$7.95. Song birds: the language of song. 2001. By S.A. Johnson. Carolrhoda, Minneapolis. 48 pp., illus. U.S.$23.95. Spiny sea stars. 2001. By C. Zuchora-Walske. Lerner, Minneapolis. 32 pp., illus. U.S.$21.95. Sure-to-win science fair projects. 2001. By J. Rhatigan. Lark Books, New York. 128 pp., illus. U.S.$21.95. *Assigned for review +Available for review. News and Comment Sea Wind: Bulletin of Ocean Voice International 5(1/2) December 2001 This special commemorative edition combines two issues between its covers and is edited by Marcia Campbell and Noel Alfonso. Sea Wind 15(1): 1-37, was the last issue Don E. McAllister was working on before his death 17 June 2001, and contains: A special notice from the editors; Fire Shrimp successfully cultured in Sri Lanka; A scientific consensus on marine reserves; Filipino fisher-folk fighting to benefit from marine biological riches; Free trade and sovereignty to protect rights of coastal communities & fish- ers; Satellite data helps save coral reefs; Selective fishing recommendations in Canada; Mapping life in the world's oceans; Ocean Voice International 2001 Annual General Meeting; Ocean Voice International Annual Report; Sea News. Sea Wind 15(2): 38-64, is dedicated to Don McAllister and contains: Memorial cover page "In memory of Don" a tableau of scleractinian corals by Roelof Idema; Ode to Grampa; Friends write in memory of Don; Dedication article: model citizen of the world; Curatorial and research contributions - Museum of Nature; Dedication article: Is bottom trawling scorched-earth fishing; Kid's korner; Tatay Don McAllister: Haribon Foundation's empowering partner; Ocean Voice International member- ship form. For membership rates contact Ocean Voice Inter- national, Box 37026, 3332 McCarthy Road, Ottawa, Ontario K1V OW0O; Telephone (613) 721-4541; Fax (613) 721-4562; Office Manager at e-mail: ocean- voice.superaje.com; home page: http://www.ovi.ca. Biodiversity: Journal of Life on Earth 3(1) February 2002 Contents: Teaming up to conserve the biodiversity of Western Kenys (Eusebius J. Mukhwana) — Some reflec- tions on the work of Don E. McAllister (Ted Mosquin) — Marine biodiversity of of Canada: Threats and conservation solutions — Black widow spiders: An outline of diversity (Charles D. Dondale) — The Giant Jamaican Swallowtail (Papilio homerus F.) — Going, going... (Peter W. Hall) — Forum: Omission of Biodiversity measurement and human population issues - shocking! (Don Kerr) -- Editor's Corner (Ted Mosquin) — Species by species [Chocolate, Theobroma cacao] — Biodiversity News — Book Reviews — Announcements. The new editor of Biodiversity is Dr. Ted Mosquin; Catherine Ripley remains as Managing Editor. Biodiversity is published by The Tropical Conservancy, 94 Four Seasons Drive, Ottawa, Ontario K2E 7S1 Canada, e-mail: tropical @synapse.net; URL: http://www.synapse.net/~tropical The Boreal Dip Net: Newsletter of the Canadian Amphibian and Reptile Conservation Network 6(1) December 2001 This issue highlights the CARCNET meetings held in Prince Edward Island in 18-22 October 2001. [List of abstracts from the meeting is available on the website: http:\www.carcnet.ca/]. Contents: Editor's note [Kerrie Serben]; Report from the Chair [Christine Bishop]; CAR- CNET Resolution to help marine trutles in Canada; 2001 CARCNET Award Winners: The Blue Racer Award [Francis R. Cook]; Silver Salamander Awards [Nature Trust of New Brunswick for establishment of Hyla Park (Fredericton) and Ducks Unlimited for their Small Marsh Program]; DigitalFrog International/CARCN Scholarship [Shana Truant: A comparison of four different salamander sampling techniques and a study of salamander popula- tions that reside in riparian reserves in Ontario's mixed- wood forests]; Student Platform presentations; Abstracts/Lecture notes from Keynote Speakers: Sherman Bleakney, Francis R. Cook, Michael C. James; Field trip - Revisiting Old Study Sites in PEI [Shana Truant]; The rare and secretive Sharp-tailed Snake [David Cunnington]; Fracine the Poster Frog ["Its not easy being green": Kejimkujik National Park: Peter Hope]; CARCNET List of Directors for 2001-2002. Recovery: An Endangered Species Newsletter (19) January 2002 Published by the Canadian Wildlife Servis this issue contains: Recovery Highlights: Tracking turtles; Protecting habitat; Taking flight; Saving Ontario's mussels [David Wylynko]; Special Report: Endangered Species Recovery Fund: Supporting a diverse array of projects; Bowhead whale strategy releasd: Long-term conservation, ecosystem approach planned for eastern arctic population; Wood Bison recovery plan approved; Scientists launch beluga recovery; Caribou committee proposed; Recovering Newfoundland plants; Updates: Workshop held; Report released; Assessing soecies; Monitoring migrations: Ham operators assist scientists in search for wild species in migration; Researchers map biodiversity; New findings in swan research; New publications; Awards; Site seeing; Featured Species: Rador reveals bird's secretive behaviour {Marbled Murrelet]. Contact: Recovery, Canadian Wildlife Service, Environment Canada, Ottawa, Ontrio KIA OH3. Web site: www.cws-scf.ec.gc/recovery/archive.html 177 178 THE CANADIAN FIELD-NATURALIST Vol. 116 RENEW: National Recovery Plan Number 21: The Wood Bison, Bison bison athabascae The Recovery of Nationally Endangered Wildlife [Canada] has issued its "National Recovery Plan for the Wood Bison, Bison bison athabascae as Plan Number 21, October 2001. 50 pages. It was prepared by a team consist- ing of C. Cormack Gates, University of Calgary; Robert O. Stephenson, Alaska Department of Fish and Game; Hal W. Reynolds, Canadian Wildlife Service, Environment Canada; C. G. Van Zyll de Jong [deceased May 1997, for- merly mammalogist, Canadian Museum of Nature]; Helen Schwantje, British Columbia Ministry of Environment, Lands and Parks; Manfred Hoefs, Yukon Department of Renewable Resources; John Nishi, Northwest Territories, Wildlife and Economic Development; Normand Cool and Jane Chisholm, Parks Canada Agency; Adam James, Alberta Fish and Wildlife Service, Sustainable Resource Development; Bill Koonz, Manitoba Department of Natural Resources. Copies are available from Recovery Secretariat, c/o Canadian Wildlife Service, Environment Canada, Ottawa, Ontario K1A 0H3; Tel 819-953-1410; Fax 819- 994-3684. Alberta Wildlife Status Reports numbers 37, 38, 39 The Fisheries and Wildlife Management Division of the Alberta Natural Resource Status and Assessment Branch, Alberta Environmental Protection, has released new Wildlife Status Reports. The Series Editor is Sherry Frazer (37), Sherry Frazer and Robin Gutsell (38 and 39); the Senior Editor for (37) is Isabelle M. G. Michaud; and the illustrations are by Brian Huffman for all three. For a list- ing earlier numbers in the series, see The Canadian Field- Naturalist 112(1): 169 for 1-11; 113(2): 311 for 12-17; 113(4): 686 for 18-21; 114(1): 151 for 22-25; 115(2): 390 for 26-31; 115(3): 000 for 32-36. Recent reports issued in 2001 are: 37. Status of the Grizly Bear (Ursus arctos) in Alberta, by John L. Kansas. 43 pages. 38. Status of the Wood Bison (Bison bison athabascae) in Alberta, by Jonathan A. Mitchell and C. Cormick Gates. 32 pages. 39. Status of the Bull Trout (Salvelinus confluentus) in Alberta, by John R. Post and Fiona D. Johnson. 40 pages. For copies contact the Information Centre - Publications Alberta Environment/Alberta Sustainable Resource Development, Fish and Wildlife Division, Main Floor, Great West Life Building, 9920 — 108 Street, Edmonton, Alberta T5K 2M4 Canada (telephone: (780) 422-2079), OR Information Service Alberta Environment/Sustainable Resource Development, #100, 3115 — 12 Street NE, Calgary, Alberta T2E 7J2, Canada (telephone: (403) 297- 3362); web site: http://www3.gov.ab.ca/srd/fw/status/ index.html. Ontario Natural Heritage Information Centre Newsletter 6(1) Winter 2002 Contents of this 20 page issue: Great Lakes Ecoregional Planning Project — NHIC Prepares Ontario's first ‘““General Status of Wild Species” report — NHIC Assists with the Georgian Bay Coast Project — Kawartha Highlands Update — Zoology: Species at Risk Biologists Assist with Surveys of Threatened Fish Species — NHIC Hosts the First Great Lakes Odonata Meeting — Botany: 2001 Botanical Highlights — Community Ecology: Rare Vegetation of Ontario: Database Cliffs of Northwestern Ontario — News and Notes: NHIC Establishes New Partner Agreement — NHIC Prepares Report on the Occurrence of Species at Risk in OMNR Districts — Trent Marine Turtle Newsletter (95) The January 2002 issue, 36 pages, contains: GUEST EpITORIAL: The swampland of sea turtle conservation: in search of a philosophy — ARTICLEs: Status of the sea turtle trade in Alexandria's fish market — Direct carapacial attachment of satellite using orthopedic bioabsorbable mini-anchor screwws on Leatherback Turtles in Culebra, Puerto Rico — Notes: The reproductive status of marine turtles nesting in the Cayman Islands: work in progress — occurrence terminology for marine turtles — sea turtle research and conservation project in Cipara, Paria University-NHIC Internships in Conservation Biology — NHiC Participates in First Pelee Island Endangered Species Festival — Updates to the OMNR and COSEWIC Lists of Species at Risk — “Bioconservation and Systematics” Proceedings Now Available — Farewell to Jarmo Jalava — Book Reviews — Focus on Gordon Wichert — Greetings from NIHC's New Coordinator — NHIC Staff List. Mailing address for Natural Heritage Information Centre: 300 Water Street, 2nd Floor, North Tower, P.O. Box 7000, Peterborough, Ontario K9J 8M5, Canada. Web page: http://www.mnr.gov.on.ca/MNR/nhic.html Peninsula, Sucre State, Venezuela: Preliminary results of the 2000 nesting season — Project update: University Progject for the study of conservation of Cuban sea turtles: completion of year 3 — ANNOUNCEMENTS — BOOK REVIEWS — News & LEGAL BRIEFS — RECENT PUBLICATIONS. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Research Group, School of Biological Sciences, University of Wales, Swansea, SA2 8PP Wales, United Kingdom; e-mail 2002 MTN @swan.ac.uk; Fax +44 1792 295447. Subscriptions to the MTN and donations towards the production of MTN and its Spanish edition NTM [Noticiero de Tortugas Marinas] should be sent to Marine Turtle Newsletter c/o NEWS AND COMMENT 179 Chelonian Research Foundation, 168 Goodrich Street, Lunenburg, Massachusetts 01462 USA; e-mail RhodinCRF@aol.com; fax + 1 978 582 6279. MTN web- site is: Froglog: Newsletter of the Declining Amphibian Populations Task Force (49) Number 49, February 2002, contains: Crisis less severe for Po Valley Spadefoot Pelobates fuscus insubricus (Vincenzo Ferri Italian DAPTF Working Group & Toads Project) — Effects of Habitat Disturbance on a Frog Community in a Mexican Dry Forest (Ireri Suazo-Ortuno) — Amphibian Monitoring in Africa (Cote d'Ivoire, Kenya) and Asia (Borneo) (from Global Amphbian Diversity Anaysis Group) — Amphibian decline in the Kariba Wilderness Area, Zimbabwe (Peter Taylor) — Froglog Shorts — Publications of Interest. Froglog is the bi-monthly newsletter of the Declining Amphibian Populations Task Force of The World Conservation Union (IUCN)/Species Survival Commission (SSC) and is supported by The Open University, The World Congress of Herpetology, The Smithsonian Institution, and Harvard University. The newsletter is Edited by John W. Wilkinson, Department of Biological Sciences, The Open University, Walton Hall, Milton Keynes, MK7 6AA, United Kingdom; e-mail: daptf@ open.ac.uk. Funding for Froglog is underwritten by the Detroit Zoological Institute, P. O. Box 39, Michigan 48068-0039, USA. The 124th Annual Business Meeting of The Ottawa Field-Naturalists’ Club: 7 January 2003 The 124th Annual Business Meeting of The Ottawa Field-Naturalists' Club will be held in the auditorium of Victoria Memorial Museum (Canadian Museum of Nature), McLeod and Metcalfe streets, Ottawa, on Tuesday 7 January 2003 at 7:30 p.m. (19:30 h). The Council for 2003 will be elected at this meeting and a brief review of the activities during 2002 will be given, as well as a statement of the Club's finances. KEN ALLISON Recording Secretary Call for Nominations: The Ottawa Field-Naturalist's Club 2003 Council Candidates for Council may be nominated by any mem- ber of The Ottawa Field-Naturalist's Club. Nominations require the signature of the nominator and a statement of willingness to serve in the position for which nominated by the nominee. Some relevant background information on the nominee should also be provided. FENJA BRODO Chair, Nominating Committee Call for Nominations: The Ottawa Field-Naturalists’ Club 2002 Awards Nominations are requested from members of The Ottawa Field-Naturalists' Club for the following: Honorary Membership, Member of the Year, George McGee Service Award, Conservation Award: Member, Conservation Award: Non-member, and the Anne Hanes Natural History Award. Descriptions of these awards have appeared in The Canadian Field-Naturalist 113(4): 689. With the exception of nominations for Honorary Member and Conservation Award: Non-member, all nominees must be members of The Ottawa Field-Naturalist Club in good standing. IRVIN BRODO Chair, Awards Committee 180 Letters to the Editor: THE CANADIAN FIELD-NATURALIST Vol. 116 Blastomycosis in Free Ranging Wolves, Canis lupus, on the North Shore of Lake Superior, Ontario — A response to P. Krizan The July-September 2000 issue of the Canadian Field-Naturalist 114(3): 491-493 contained an article titled “Blastomycosis in a Free Ranging Wolf, Canis lupus, on the North Shore of Lake Superior, Ontario”, authored by Mr. Peter Krizan. We regret that Mr. Krizan published the article without institutional or contractual approval of the researchers who collabo- rated in the collection, analyses, and interpretation of the data. We view the independent publication and appropriation of these results as an unprofessional assumption of the authorship and very serious breach of professional ethics. For the record, the research was conducted under federal and provincial permits issued to Dr. Paul Paquet as primary investigator. By com- mon agreement that included Mr. Krizan; the study team had assigned senior authorship to Dr. Doug Campbell of OVC, Pathobiology, Ontario Region Wildlife Health Centre, University of Guelph. In alphabetical order, coauthors were Frank Burrows (Parks Canada), Anne Forshner (University of Alberta), Peter Krizan (Acadia University), Graham Neale (University of Montana), Paul Paquet (University of Calgary), and Keith Wade (Parks Canada). The information included in Mr. Krizan’s initial report is accurate but incomplete. We take this opportunity to expand the original note. Blasto- mycosis is a chronic fungal disease that affects the pulmonary system of humans and dogs, and occa- sionally other animals (Jungerman and Schwartzman 1972; Stroud and Coles 1980; Legendre et al. 1981; Thiel et al. 1995). The fungus is enzootic in Min- nesota (Schlosser 1980) and Wisconsin (Sarosi et al. 1979; McDonough and Kuzman 1980) but until now, had not been reported in other North American wolf populations. From 1995-1999, three adult radiocol- lared wolves from our study sample (n = 26) were diagnosed with Blastomycosis at the time of death, though only one (Krizan 2000) was determined to have died of the disease. One wolf (Spirit, male) was shot and another (Moon, female) killed by other wolves (Forshner 2000). The latter was in extremely poor condition and evidence at the site of her death suggested pack mates killed her. All three wolves occupied home ranges north of Pukaskwa National Park and were members of different packs. Recently, these wolves were shown to be genetically distinct from the Gray Wolves (C. lupus) occurring within Pukaskwa National Park (Wilson et al. 2000). Both types of wolves are known to occur in Minnesota and Wisconsin but the original reports of blastomy- cosis infections in these regions preceded genetic typing. Consequently, blastomycosis has not been unambiguously confirmed in both types of wolves. Literature Cited Forshner, S. A. 2000. Population dynamics and limitation of wolves (Canis lupus) in the Greater Pukaskwa Ecosystem, Ontario. M.Sc thesis, University of Alberta, Edmonton, Alberta, 129 pages. Jungerman, P. F., and R. M. Schwartzman. 1972. Veter- inary medical mycology. Lea and Febiger, Philadelphia, Pennsylvania, USA. 200 pages. Legendre, A. M., B. A. Selcer, D. F. Edwards, and R. Stevens. 1984. Treatment of canine blastomycosis with amphotericin B and ketoconazole. Journal of the Ameri- can Veterinary Medical Association 184: 1249-1254. McDonough, E. S., and J. F. Kuzma. 1980. Epide- miological studies on blastomycosis in the state of Wisconsin. Sabouraudia 18: 1373-183. Sarosi, G. A., M. R. Eckman, S. F. Davies, and W. K. Laskey. 1979. Canine blastomycosis as a harbinger of human disease. Annals of Internal Medicine 91: 773-735. Stroud, R. K., and E. M. Coles. 1980. Blastomycosis in an African lion. Journal of the American Veterinary Medical Association 177: 842-844. Thiel, R. P., L. D. Mech, G. R. Ruth, J. R. Archer, and L. Kaufman. 1987. Blastomycosis in wild wolves. Journal of Wildlife Disease 23: 321-323. Wilson, P. J., S. Grewal, I. D. Lawford, J. Heal, A. G. Granacki, D. Pennock, J. B. Theberge, M. T. Theberge, D. Voigt, W. Waddell, R. E. Chambers, P. C. Paquet, G. Goulet, D. Cluff, and B. N. White. 2000. DNA profiles of the eastern Canadian wolf and the red wolf provide evidence for a common evolution- ary history independent of the gray wolf. Canadian Journal of Zoology 78: 2156-2166. PAUL C. PAQUET (Conservation Science Inc. and University of Calgary)! FRANK Burrows (Parks Canada | and Lakehead University) ANNE FORSHNER (University of Alberta) GRAHAM NEALE (University of Montana) | KEITH WADE (Parks Canada) | Received 12 May 2001 'Box 150, Meacham, Saskatchewan SOK 2V0 Canada; | email: ppaquet @sk.sympatico,ca 2002 NEWS AND COMMENT 181 Blastomycosis in a Free ranging Wolf, Canis lupus, on the North Shore of Lake Superior, Ontario — A reply to P. Paquet et al. Paquet et al. report interesting results that add valuable information to the original note by Krizan 2000 and suggest that Blastomycosis is more preva- lent than was originally thought and may not always be fatal. My note (Krizan 2000) was one finding of my Masters thesis that was started in 1993 and was completed and defended in 1997 (Krizan 1997). It should be noted that the additional data provided by Paquet et al. were collected after the completion of my thesis. In both publications (Krizan 2000, 1997), I acknowledged the members and agency involved in the co-operative project for their role and participa- tion. Parks Canada (Pukaskwa National Park) sup- ported the Pukaskwa National Park Predator Prey Process Project (P5) through logistical support. In addition, many volunteers and other researchers working on different aspects of this project were involved to different degrees; those who had con- tributed directly to the collection of data were men- tioned by name in the acknowledgment sections of Krizan 1997 and 2000. I once again acknowledge the role of some of the co-authors of Paquet et al. in the overall project and their recent contribution. I was one of three students completing a Masters thesis from 1993 to 1997. The only agreement that I had with Parks Canada and/or P. Paquet, who was the principal investigator contracted by Parks 'Canada was that Parks Canada would supply the logistical support for my project and in return, I ‘would collect, analyze and publish my results and make my thesis and raw data available to the 'Pukaskwa National Park office. I have fulfilled all of those responsibilities. During 1993 to 1997 I was an ‘independent student with no contractual agreements with any of the co-authors of Paquet et al. I was the ‘only one that analyzed data that was used in the Krizan 2000 publication except for the necropsy that ‘was done by Dr. Doug Campbell (as was acknowl- edged). I had discussed Dr. Campbell’s necropsy iresults with him and he provided me with relevant literature. To the best of my knowledge, the research results of the Masters thesis are owned by copyright and are not subject to institutional or contractual consent. My research interest was to describe the travel patterns of Wolves and how these movements were affected by human presence, activity, and land- scape differences in and outside of Pukaskwa National Park. The case of blastomycosis was inter- esting in that infection by this fungus was probably due to the fact that this Wolf was a lone animal that exposed itself to areas by travelling in environments that may have had a higher probability of exposure to the fungal spores. Also interesting was the fact that the travel distances between locations dimin- ished as the disease affected the ability of the Wolf to travel. To the best of my knowledge, I was the only person on the P5 project team that was interest- ed in the spatial use of Wolves. As with any project, there are many people that are involved to various degrees and I’ve acknowledged those who had con- tributed directly to the collection of data. Two of the co-authors of Paquet et al. were also students working on their theses of various but dif- ferent topics. I am confused about the role and co- authorship of Miss Forshner. I met Miss Forshner once as I was leaving the project. I had never worked with her nor did I ever discuss authorship of publica- tions. The implied consent to co-author any publica- tion in the order that Paquet et al. suggest is incor- rect. Literature Cited Krizan, P. 1997. The effects of human land development, landscape characteristics, and prey density on the spatial distribution of Wolves (Canis lupus) on the north shore of Lake Superior, Ontario. M.Sc. thesis, Acadia Univer- sity, Wolfville, Nova Scotia. 108 pages. Krizan, P. 2000. Blastomycosis in a free ranging lone Wolf, Canis lupus, on the north shore of Lake Superior, Ontario. Canadian Field-Naturalist 114: 491-493. PETER KRIZAN P.O. Box 1167 Iqaluit, Nunavut XOA OHO Canada Received 5 December 2001 - Advice for Contributors to The Canadian Field-Naturalist Content The Canadian Field-Naturalist is a medium for the publi- cation of scientific papers by amateur and professional natu- ralists or field-biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All readers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and com- ment items if judged by the Editor to be of interest to read- ers and subscribers, and book reviews. Please correspond with the Book Review Editor concerning suitability of manuscripts for this section. For further information consult: A Publication Policy for the Ottawa Field-Naturalists’ Club, 1983. The Canadian Field-Naturalist 97(2): 231-234. 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Reprints An order form for the purchase of reprints will accom- pany the galley proofs sent to the authors. FRANCIS R. CooK, Editor RR 3 North Augusta, Ontario KOG 1RO, Canada 182 TABLE OF CONTENTS (concluded) /olf, Canis lupus, response to domestic sled dog, Canis familiaris, activities in central Yukon GERALD W. KUZyYK and KRISTIN M. KUZYK irst record of an Eastern Coyote, Canis latrans, in Labrador Tony E. CHUBBs and FRANK R. PHILLIPS ong distance movement by a Coyote, Canis latrans, and a Red Fox, Vulpes vulpes, in Ontario: Implications for disease-spread RICHARD C. ROSATTE redation of Wolves, Canis lupus, on Wolverines, Gulo gulo,and an American Marten, Martes americana, in Alaska KEVIN S. WHITE, HOWARD N. GOLDEN, Kris J. HUNDERTMARK, and GERALD R. LEE feek’s Halfbeak, Hyporhamphus meeki, and Flying Gurnard, Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia A. JAMIE F. GIBSON and RANSOM A. MYERS pparent capture myopathy in Hoary Bats, Lasiurus cinereus: A cautionary note THOMAS S. JUNG, IAN D. THoMPsoN, M. BRIAN C. HICKEY, and ROGER D. TITMAN lobbing Black-billed Magpie, Pica hudsonia, killed by Cooper’s Hawk, Accipiter cooperii GEOFFREY L. HOLROYD reeding season of Wolves, Canis lupus, in relation to latitude L. DAviD MECH fistorical Feature Article anada and the “buffalo”, Bison bison: A tale of two herds W. A. FULLER 00k Reviews oology: Birds, Mammals & Reptiles of the Galapagos Islands — Wisconsin Fishes 2000: Status and Distribution — Marine Mammals of the Pacific Northwest: A Concise and Comprehensive Waterproof Guide — Manitoba Birds — Heron Conservation — The Field Guide to Birds of Australia — Katydids and Bush-Crickets: Reproductive Behavior and Evolution of the Tettigoniideae — A Manual for Wildlife Radio Tagging — Bird Census Techniques — Flying Foxes: Fruit and Blossom Bats of Australia otany: Guide to Standard Floras of the World: An Annotated Geographically Arranged Systematic Bibliography of the Principal Floras, Enumerations, Checklists, and Chorological Atlases of Different Areas, Second Edition nvironment: Their Father’s Work: Casting Nets with the World’s Fishermen — AAAS Atlas of Population and Environment — Making Better Environmental Decisions — Scientific Method for Ecological Research liscellaneous: The Last Great Sea: A Voyage through Human and Natural History of the North Pacific Ocean ew Titles lews and Comments: 2a Wind: Bulletin of Ocean Voice International 5(1/2) December 2001 — Biodiversity: Journal of Life / on Earth 3(1) February 2002 — The Boreal Dip Net: Newsletter of the Canadian Amphibian and Reptile Conservation Network 6(1) December 2001 — Recovery: An Endangered Species _ Newsletter (19) January 2002 —RENEW: National Recovery Plan Number 21: The Wood Bison, _ Bison bison athabascae — Alberta Wildlife Status Reports numbers 37, 38, 39 — Ontario Natural | Heritage Information Centre Newsletter 6(1) Winter 2002 — Marine Turtle Newsletter (95) — Frogleg: Newsletter of the Dectining Amphibian Population Task Forces (49) — Notice of the 124th Annual Business Meeting of The Ottawa Field-Naturalists’ Club: 7 January 2003 — Call for Nominations: The Ottawa Field-Naturalist’s Club 2003 Council — Call for Nominations: The Ottawa Field-Naturalists’ Club 2002 Awards ‘etters to the Editor: Blastomycosis in free-ranging Wolves, Canis lupus, on the north shore of Lake _ Superior, Ontario: A response to P. Krizan; A reply to Paquet et al. dvice to Contributors lailing date of the previous issue 115(4): 9 August 2002 ve 125 127 129 132 134 136 137 139 141 160 168 169 172 174 THE CANADIAN FIELD-NATURALIST : Volume 116, Number 1 a Articles Spread and disappearance of the Greater Prairie-Chicken, Tympanuchus cupido, on the Canadian prairies and adjacent areas C. STUART HOUSTON Status of Common Eiders, Somateria mollissima, nesting in the Digges Sound region, Nunavut J. MARK HIPFNER, H. GRANT GILCHRIST, ANTHONY J. GASTON, and DAvID K. CAIRNS Flathead Chubs, Platygobio gracilis, in the upper Missouri River: The biology of a species at risk in an endangered habitat oo a ~ SHANNON J. FISHER, DAVID W. WILLIS, See * = MICHAEL M. OLson, and STEVEN C. KRENTZ Abundance and distribution of breeding waterfowl in the Great Clay Belt of northern Ontario _R. KENYON Ross, KENNETH F. ABRAHAM, TED R. GADAWSKI, ROBERT S. REMPEL, T. SHANE GABOR, and RON MAHER Resilience of Foothills Rough Fescue, Festuca campestris, rangeland to wildfire EDWARD W. Bork, BARRY W. ADAMS, and WALTER D. WILLMS Survival, fates, and success of SED Beavers, Castor canadensis, in Wyoming Mark C. McKINstTRY and STANLEY H. ANDERSON Songbird community composition versus forest rotation age in Saskatchewan boreal mixedwood forest ENID E. CUMMING and ANTONY W. DIAMOND Status of Redside Dace, Clinostomus elongatus, in the Lynde Creek and Pringle Creek watersheds of Lake Ontario JEFF J. ANDERSON Aquatic leaves and regeneration of last year’s staw in the arctic grass, Arctophila fulva SUSAN G. AIKEN and ROSEMARY A. BUCK Records of Northern Mockingbird, Mimus polyglottos, occurrences in North Dakota during the twentieth century LAWRENCE D. IGL and RON E. MARTIN Diets of Northern Flying Squirrels, Glaucomys sabrinus, in southeast Alaska SANJAY PYARE, WINSTON P. SMITH, JEFFREY V. NICHOLLS, and JOSEPH A. COOK Nesting activities of an Eastern Spiny Softshell Turtle, Apalone spinifera CLAUDE DAIGLE, PATRICK GALOIS, and YVES CHAGNON Long-distance movements by female White-footed Mice, Peromyscus lecopus, in extensive mixed-wood forest THOMAS J. MAIER Recent trends in stem numbers in Goldenseal, Hydrastis canadensis, populations at the northern limit of its range ADRIANNE SINCLAIR and PAUL M. CATLING A comparison of techniques for assessing amphibian assemblages on streams in the western boreal forest CYNTHIA A. PASZOWSKI, GARRY SCRIMGEOUR, BEVERLY A. GINGRAS, and SHARON KENDALL Notes A rare leucistic Spiny Dogfish, Squalus acanthias, from the Bay of Fundy, Nova Scotia BRIAN W. CoaD and JOHN GILHEN Anomalies in incisor wear of American Elk, Cervus elaphus, in the French River delta, Ontario J. HAMR, F. F. MALLory, I. A. FILION, G. S. BRowN, and M. A. Jost First record of the Hoary Bat, Lasiurus cinereus (Chiroptera: Vespertilionidae), from Prince Edward Island DONALD F. MCALPINE, FRANCES MULDOON, and ALEXANDER I. WANDELER SMITH STITUTION L NO ISSN 0008-3550 3 9088 ll 614 j 3 The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada /olume 116, Number 2 April-June 2002 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Her Excellency The Right Honourable Adrienne Clarkson, C.C., C.M.M., C.D. Governor General of Canada His Excellency John Ralston Saul, C.C. 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 infor- — mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain- ing or restoring environments of high quality for living things. Honorary Members . Edward L. Bousfield Bruce Di Labio George F. Ledingham E. Franklin Pope Donald M. Britton R. Yorke Edwards John A. Livingston William O. Pruitt, Jr. Irwin M. Brodo Anthony J. Erskine Stewart D. MacDonald Joyce and Allan Reddoc William J. Cody John M. Gillett Hue N. MacKenzie Mary E. Stuart Francis R. Cook W. Earl Godfrey Theodore Mosquin John B. Theberge Ellaine Dickson C. Stuart Houston Eugene G. Munroe Sheila Thomson Robert W. Nero \ 2002 Council President: Eleanor Zurbrigg Ronald E. Bedford Francis R. Cook David W. Moore . : Rosanne Bishop Barbara Gaertner Robert Roach Vice-President: Gary McNult ee Ee icin sind Irwin Brodo Diane Lepage Stanley Rosenbaum , John Cameron Diane Holms Louise Schwartz Recording Secretary: Ken Allison William J. Cody David Hobden David Smythe Treasurer: Frank Pope Kathy Conlan Beverly McBride Dorothy Whyte To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, Box P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net. For information on Club activities telephone (613) 722-3050 or check http//www.achilles.net/ofnc/index.htm The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. We acknowledge the financial support of the Government of Canada toward our mailing cost through the Publication Assistance Program (PAP), Heritage number 09477. Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1RO; (613) 269-3211; e-mail: feook @achilles.net Copy Editor: Elizabeth Morton Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 759-1374 Book Review Editor: Dr. J. Wilson Eedy, 9074-32 Side Road, R.R.1, Glencairn, Ontario LOM K00 e-mail: terfa@ geoniger.com Associate Editors: Robert R. Anderson RobertR. Campbell Brian W. Coad W. Earl Godfrey Donald F. McAlpine Charles D. Bird Paul M. Catling Anthony J. Erskine William O. Pruitt, Jr. | David Nagorsen Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication should be addressed to the Editor and sent by postal mail (no courier, no post requiring signature on delivery). Exception: book reviews should go directly to Book Review Editor. Subscriptions and Membership Subscription rates for individuals are $28 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $28 (individual) $30 (family) $50 (sustaining) and $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and mem- bers (including USA) must add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at $28 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications Mail Agreement number 40012317. Return Postage Guaranteay Date of this issue: April-June 2002 (February 2003). Cover: Logan Glacier and the St. Elias Mountains in Wrangell-St.Elias National Park and Preserve, Alaska. The site in the foreground is the locality of Arabis calderi G. A. Mulligan, which is new to Alaska, and also the collection locality of the mustards Braya glabella Richardson ssp. glabella, Draba cinera Adams, D. incerta Payson, D. macounii O. E. Schultz, D. oligosperma Hook. and Lesquerella arctica (Wormsk.) S. Wats. ssp. arctica, all of which were range extensions at this site. Silene menziesii Hook., Carex filifolia Nutt., and Danthonia intermedia Vasey were other range extensions from this locality. See article on notable vascular plants form Alaska by Mary B. Cook and Carl A. Roland, pages 192-304. The Canadian Field-Naturalist Volume 116, Number 2 April-June 2002 PAUL F. JONEs,! and ROBERT J. HUDSON Department of Renewable Resources, University of Alberta, Edmonton, Alberta T6G *, Canada ‘Present address: Alberta Conservation Association, 2nd Floor, YPM ‘Place, 530 — 8th Street S., Lethbridge, Alberta T1J 2J8 Canada; E-mail: paul.jones @ gov.ab.ca Jones, Paul F., and Robert J. Hudson. 2002. Winter habitat selection at three spatial scales by American Elk, Cervus ela- phus, in west-central Alberta. Canadian Field-Naturalist 116(2): 183-191. Selection by American Elk (Cervus elaphus) at the landscape (2 order), stand (3" order), and site level (4 order) was examined in west-central Alberta from 1 December 1994 to 21 March 1995. At the landscape level, elk home ranges had a lower mean road density, but greater mean unimproved access density. Elk home ranges had smaller mean patch sizes and greater patch density. There was no significant selection on the basis of food and cover composition. At a stand level, elk used the grass/meadow habitat more than expected while all other habitat types were used in proportion to their availabili- ty. At a site scale, we attempted to distinguish habitat use for feeding and bedding. Feeding sites had a lower mean percent canopy closure, percent shrub cover, stem density and tree height, but a higher mean percent grass cover than bedding sites. Feeding sites were significantly closer to unimproved access than were bedding sites. Feeding sites had more grass cover and significantly less canopy closure, percent shrub cover, spruce, pine, and fir in the canopy, and lower tree height and stem density than the control plots. They were located significantly closer to unimproved access and farther from hid- ing cover. Bedding sites had a higher percent grass cover, less spruce, pine and fir in the canopy, and a lower stem density than the control plots. In terms of spatial attributes, elk exhibited no selection when choosing bedding sites. The need to examine habitat selection patterns at more than one scale is discussed Key words: Cervus elaphus, American Elk, habitat, landscape, site, spatial scale, stand, Alberta. Resource selection is an adaptive response of ani- mals attempting to meet their life requirements and ultimately to increase fitness (Johnson 1980; Thomas and Taylor 1990; Manly et al. 1993). Choice of habi- _ tat is governed by a variety of factors including ener- gy requirements, thermal conditions, reproductive requirements, and intra- and interspecific competi- tion (Litvaitis et al. 1986; Morrison et al. 1992). Johnson (1980) defined four levels or scales of habitat selection: (1) selection of a physical or geo- graphical range, (2) selection of a home range within that geographical range, called landscape selection, (3) selection of habitat components within the home range, termed stand selection and (4) selection of food items at a particular feeding site. Lofroth (1993) expanded Johnson’s (1980) fourth-order selection to include habitat components required for such activities as mating, calving, escaping predators and for shelter. We define this level of selection as site selection. There are several possible outcomes to habitat studies conducted at multiple scales. Selection main- ly at the landscape level is important in several species such as the Spotted Owl (Strix occidentalis). Hansen et al. (1993) concluded that landscape level habitat features describing old growth forests were the best predictors of spotted owl abundance. At the other end of the spectrum, a species may select habi- tat characteristics mainly at a site level. This may be the case for generalists who are able to utilize differ- ent local conditions found in a variety of landscapes (Pearson 1993). The probable outcome of multiple level habitat selection is a species selecting habitat characteristics at more than one scale. Lofroth (1993) determined that Marten (Martes americana) selected mature to old growth seral stage habitats within their home range (landscape level), while showing no preference for any habitat type, but avoided young seral stages, xeric habitat types and wetlands (stand level). At the site level, Marten selected structural characteristics that were different from the prevailing characteristics of the habitat type (Lofroth 1993). Selection at more than one scale may be complementary to each other or selection at one level may explain why it occurred at a higher level (Pedlar et al. 1997). Previous research has suggested that habitat selec- tion by elk (Cervus elaphus) has been governed by 183 184 forage, thermal cover and hiding cover requirements (Hersey and Leege 1976; Lyon 1979; Gates and Hudson 1981; Irwin and Peek 1983; Marcum and Scott 1985; Edge et al. 1987; McCorquodale 1987). These studies have examined selection at one specif- ic level. We examined resource use by elk at the landscape (2" order), stand (3™ order) and site level (4 order). The central hypothesis was that elk select habitats for foraging, thermal cover and hiding cover at more than one scale. Our first objective was to determine if elk selected home ranges based on habi- tat components. Our second objective was to deter- mine if elk selected specific habitat types within their home ranges. Our third objective was to deter- mine whether elk selected habitat characteristics to meet different life requisites (i.e., feeding and bed- ding), and whether or not elk select specific sites within the immediately available area on the basis of non-spatial and spatial attributes. Study Area The study was conducted in west-central Alberta along the north and south facing slopes of the Athabasca River Valley near Hinton, Alberta (53°25 N, 117°35'’W). The area falls within the Lower Foothills Natural Subregion and rises in ele- vation from 500m to 1150m (Beckingham et al. 1996). The winter climate of the Lower Foothills Subregion is prone to warming by chinook winds (Beckingham et al. 1996). Mean monthly tempera- ture in winter (November through February) is -7.8°C with a total annual precipitation of 60 mm (Beckingham et al. 1996). During the study (Decem- ber 1994 to March 1995) the mean monthly tempera- ture was -7.6°C, with the study area receiving 35 mm of precipitation, predominantly as snow. Vegetation in the study area was primarily mixed- wood forest, some of which were stands regenerating after clearcut logging. The area is dominated by mature mixedwood stands of Trembling Aspen (Populus tremuloides), Lodgepole Pine (Pinus contor- ta), White and Black Spruce (Picea glauca and P. mariana respectively), and Balsam Poplar (Populus balsamifera). Low-bush Cranberry (Viburnum edule), Prickly Rose (Rosa acicularis), Green Alder (Alnus crispa), Canada Buffalo-berry (Shepherdia canaden- sis), Marsh Reed Grass (Calamagrostis canadensis), and Hairy Wild Rye (Elymus innovatus) are common understorey species (Beckingham et al. 1996). A few large meadows also were present in the study area. Methods Study Animals and Site Location From June 1993 to September 1994 elk were cap- tured using up to eight collapsible clover traps (Thompson et al. 1989) baited with salt blocks or alfalfa. Traps were placed in areas of known elk use and checked twice daily. Captured elk were fitted with a Lotek LMRT-4 radio collar containing a mor- THE CANADIAN FIELD-NATURALIST Vol. 116 tality sensor and a head up/head down activity sen- sor. All animals captured were handled with care in accordance to the principles and guidelines outlined by the Canadian Council of Animal Care. Twelve animals (10 cows and 2 bulls) were tracked through- out 24-hr days during the winter of 1994/1995. Elk were located at least twice a week from 1 December 1994 to 21 March 1995, subject to field restrictions. Animals were located after 30 November 1994 to lessen the effect of hunting on habitat selection, while the 21 March 1995 date corresponded to a shift back to spring/summer range. The necessity of obtaining approximately 20 locations per animal over the winter period prohibited a complete ran- domized design. To reduce autocorrelation of suc- cessive locations (Swihart and Slade 1985), individ- ual animals were located at least 72 hours apart (mean = 95.5 hours). A Telonics TR-2 receiver with scanner and three element yagi antenna received signals from radio col- lared elk. Locations were taken from the ground from fixed bearing sites, with animal locations determined from 3-5 intersecting compass bearings. The interval between bearings was usually less than 25 minutes. We attempted to minimize distance between receiver and collared elk without disturbing the elk. We tried to visit each fix within 24 hours to confirm elk activi- ty (feeding, bedding, or travelling) at that point (84% confirmed). Additional feeding and bedding sites were determined by visual sightings of non-collared elk feeding or bedded, and the discovery of feeding or bedding sites while performing radio telemetry. Only one bed or feeding site was measured per group of animals. Universal Transverse Mercator (UTM) coordinates for all locations were determined using a Trimble Global Positioning System (GPS) unit (Trimble Navigation 1992a) and differentially cor- rected using the software Pathfinder (Trimble Navigation 1992b). Landscape Level Selection Only study animals that were located = 18 times during the winter were included for analysis at the landscape level (n = 9; 8 cows and | bull). The 95% adaptive kernal (ADK) home ranges were estimated for nine of the 12 collared elk using the program CALHOME (Kie et al. 1996). The 95% ADK UTM coordinates for the nine elk home ranges were imported into ARC/INFO (ESRI 1990) for spatial analysis. Weldwood of Canada’s (Hinton Division) forest cover (Phase III data), roads, unimproved access (seismic lines, cut-lines, power lines and pipeline right-of-ways) and forest harvest data were summarized for each home range. To determine availability of habitat at the landscape level, nine random circular home ranges equal to the mean area of the nine elk home ranges were created in ARC/INFO (ESRI 1990). We termed these random circular home ranges available home ranges (AHR). 2002 Each AHR was randomly placed but had to fall com- pletely within the study area boundary. The forest cover, roads, unimproved access, and forest harvest data were clipped and summarized for each AHR (n for available = 9). Stands were classified using the Phase III cate- gories of the Alberta Forest Inventory method (Alberta Forest Service 1981) into food, thermal cover and hiding cover separately for each home range and AHR. A stand with canopy closure no greater than 50% was classified as food. Thermal cover was a stand with canopy closure of 51% or greater and a tree height of 12 m or greater. Hiding cover was defined as a stand with canopy closure of 50% or greater. These definitions were based on standard definitions for elk habitat (Thomas et al. 1979) with modifications to represent available local habitat types. A stand could be classified into more than one type under the assumption that elk utilize certain stands for more than one purpose. For exam- ple a stand could be used for both thermal and hiding cover. For each home range and AHR, the road density, unimproved access density, patch density, and mean patch size were calculated. Road density was calcu- lated as the kilometers of permanent roads found within an elk’s home range or AHR, while unim- proved access density was calculated as the kilome- ters of seismic lines, cut lines, power lines and gas line rights-of-way found within an elk’s home range or AHR. Patch density was defined as the number of forest cover stands (> 1 ha) per km’, while mean patch size was the mean size of forest cover stands. Finally the percent of each home range and AHR previously logged (between 1957 and 1985) was determined. To evaluate habitat selection at the land- scape level, we compared the composition of the elk home ranges to the AHR using the Kruskal-Wallis test. Stand Level Selection Locations contained within the 95% ADK home range for each elk (n = 9) were classified as 1 of 10 habitat types: grass/meadow, regenerating cut, water, sapling, pole, open mature, closed mature, open old growth, closed old growth or other. Open stands had a canopy cover less than 50% while closed stands had a canopy greater than 49%. White and Garrott (1990) recommended determining habitat selection at an individual level because different animals may use the habitat differently. However, the number of relocations for each animal was inadequate for anal- ysis at an individual level (Neu et al. 1974). There- fore, locations from each collared animal were pooled. Habitat availability was determined by aver- aging the percent composition of each habitat type within each elk’s home range. A chi-square test was used to determine if elk used each habitat type in proportion to its availability (Neu et al. 1974). If the JONES AND HUDSON: WINTER HABITAT SELECTION BY AMERICAN ELK 185 null hypothesis that elk were using the habitat in proportion to its availability was rejected, a Bonferroni Z statistic was calculated for each habitat type to determine if it was used more or less than expected (Neu et al. 1974). Site Level Selection Within a home range, specific sites are used for specific purposes. For all feeding and bedding sites, a number of vegetative and spatial variables were determined. Visual estimates (percent) of grass cover, herb cover, and shrub cover were assessed in a 0.04 ha circular plot (radius 11.3 m) centered at the site location. Also within the 0.04 ha plot the num- ber of trees were counted and classified to determine percent spruce, pine and fir in the canopy and tree density (scaled to #/ha X 1000). Average tree height was determined by averaging the height of five trees, representing the stand, as determined using a cli- nometer (Bessie 1995). Canopy closure was estimat- ed at the center of the plot using a spherical den- siometer. All measurements were taken between June 1995 and September 1995, except percent can- opy closure, which was measured in May. Spatial variables were estimated from maps pro- duced by overlaying the feeding and bedding loca- tions with Weldwood of Canada’s (Hinton Division) forest cover, road and seismic line data using ARC/INFO software (ESRI 1990). Distances to major roads, minor roads, unimproved access, forage stands, thermal cover stands and hiding cover stands were determined. All variables were estimated in meters. Major roads were defined as any hard packed road, open year round and maintained, while minor roads were defined as roads that are not main- tained year round and are usually only accessible by 4x4 vehicle during the winter. Unimproved access was an inclusive term for seismic lines, cut lines, power lines and gas line right-of-ways. A l-way ANOVA was used to determine if the mean spatial and non-spatial attributes at elk feeding sites were significantly different than those found at elk bed- ding sites. To determine if elk selected particular sites for feeding and bedding, we located a paired control plot 300 m from the feeding or bedding site, in a random direction (Brown 1994). The non-spatial and spatial variables estimated at the feeding and bedding sites were also assessed at the control plots. For each non- spatial and spatial attribute, the means for the feed- ing and bedding sites were compared to the means at their paired control plot using a paired t-test. All tests were preformed using SPSS (Norusis 1993). Results Landscape Level Selection The mean 95% ADK home range size for the nine collared elk was 23 km? (range: 12-53 km’). Four landscape variables were significantly different 186 THE CANADIAN FIELD-NATURALISTE Vol. 116 Table 1. Mean attributes for elk home ranges (n = 9) and available home ranges (AHR) (n = 9) for the winter of 1994-1995 in west-central Alberta. Elk Home Range* M+ SE Habitat Variables Food (%) 49.22 + 1.93 Thermal Cover (%) A3.28 = Qf. Hiding Cover (%) 88.16 + 1.28 Harvested (%) 10,504 3.37 Density Variables Road Density (km/km*) 0.32 + 0.05 Unimproved Access Density (km/km7) 3.18 + 0.16 Patch Density(#/km?) 42.99 + 6.12 Mean Patch Size (km7) 0.03 + 0.004 AHR? Mean + SE 48.47 + 6.95 41.92 +6.17 82.21 + 4.88 7.92 + 4.10 0:82 0:22 2.48 + 0.24 21.67 + 4.85 0.06 + 0.009 P Value‘ 0.453 0.427 0.508 0.269 0.005 0.038 0.012 0.007 4 Mean home range size = 23 km? > Available home range size = 23 km? © P values are the results from Kruskal-Wallis tests. between the elk home ranges and the AHR (Table 1). Elk home ranges had a lower mean road density, but a greater mean unimproved access density than the AHR. They also had smaller mean patch size and a greater patch density. There was no significant dif- ference in the composition of the home ranges and AHR in terms of food and cover composition. Stand Level Selection For all elk home ranges combined (n = 9), the dominant cover type was closed old growth while sapling and water (1.e., lake and river) were the least prevalent cover types (Table 2). Grass/meadow accounted for 7% of the combined home ranges, while regenerating cuts accounted for 10%. The grass/meadow cover type was used more than expected, while all other habitat types were used in proportion to their availability (Table 2). Site Level Selection Five vegetative variables were significantly dif- ferent between the 96 elk feeding and 62 bedding sites (Table 3). Compared to bedding sites, feeding sites had a lower mean percent canopy closure, per- cent shrub cover, tree density and tree height, but a higher mean percent grass cover. Feeding sites were significantly closer to unimproved access than bed- ding sites (Table 3). Although this comparison high- lighted use of specific resources for specific activi- ties, we also attempted to determine selection relative to availability. All of the non-spatial habitat attributes were sig- nificantly different between the feeding sites and control plots, except for percent herbaceous cover (Table 4). Feeding sites had significantly less canopy closure, and less spruce, pine, and fir in the canopy than the control plots. Feeding sites had a signifi- cantly lower tree height and stem density than the control plots. Feeding sites had a greater percent grass cover and less percent shrub cover than the control plots. Only two of the five spatial variables were significantly different between elk feeding sites and the control plots (Table 4). Feeding sites were Table 2. Habitat use at a stand level by elk (n = 9) in west-central Alberta during the winter of 1994-1995. Observed Expected Bonferroni 95% Confidence Habitat Available (%)* (n) (n) Ae Interval Use Grass / Meadow 7 43 13 64.94 0.137 - 0.283 More Regenerating Cut 10 21 20 0.03 0.048 - 0.157 Equal Water 2 0 4 3.83 0.000 - 0.000 No Use Sapling 3 4 5 0.39 -0.005 - 0.044 Equal Pole 10 12 20 3:22 0.016 - 0.101 Equal Open Mature 7 18 16 O37 0.037 - 0.139 Equal Closed Mature 16 40 33 1.56 0.124 - 0.266 Equal Open Old-growth 1] 15 24 a7 0.026 - 0.120 Equal Closed Old-growth 26 39 53 Ce | 0.120 - 0.126 Equal Other 8 1) 17 0.85 0.020 - 0.107 Equal “Pooling all elk locations and defining the area of availability as the mean of each habitat type found within each elk’s 95% ADK home range 2002 JONES AND HUDSON: WINTER HABITAT SELECTION BY AMERICAN ELK Table 3. Mean non-spatial and spatial attributes at elk feeding (n=96) and bedding sites (n=62) in west-central Alberta during the winter of 1994-1995. Non-spatial Variables Canopy Closure (%) Grass Cover (%) Herb Cover (%) Shrub Cover (%) Spruce, Pine, & Fir in the canopy (%) Stem Density (#/ ha < 1000) Tree Height (m) Spatial Variables Major Roads Minor Roads Elk Feeding Sites Elk Bedding Sites M+SE Mean + SE P Value‘ 31.34 + 3.45 64.68 + 4.03 <0.001 5973 245 / 34.95 + 3.30 <0.001 19.00 + 1.16 16.85 + 0.89 0.185 15:15, 2149 27.37 + 1.76 <0.001 28.17 + 4.20 38.53 + 5.07 0.120 131,2°0:19 2.03 + 0.17 0.008 S57 HOTZ 13.38 + 0.86 <0.001 1093 + 66.06 1244 + 87.46 0.164 601 + 70.05 763 + 69.90 0.121 102 + 11.30 165+ 19.59 0.004 187 Unimproved Access 4 P-values are the result of 1-way ANOVA’s. located significantly closer to unimproved access and further from hiding cover than expected on the basis of availability. Bedding sites had a significantly higher percent grass cover, and significantly less spruce, pine and fir in the canopy than control plots (Table 5). Elk selected bedding sites with significantly fewer trees per ha than available. There was no significant dif- ference between bedding sites and the control plots in terms of the distance to major and minor roads, and the distance to unimproved access (Table 5). There was no significant difference between the dis- tance from the bedding site to foraging areas and the distance from the control plots to foraging areas (Table 5). Discusion The results from multi-scale habitat studies are largely dependent on the selection of variables cho- sen to investigate (Pedlar et al. 1997). Erroneous conclusions may occur if important variables are selected at one scale and frivolous variables are selected at another (Pedlar et al. 1997). Selection of variables based on the findings of previous studies may alleviate the problem of variable selection. In our study we selected variables at each scale that represented characteristics of food, thermal cover and hiding cover; requisites that have been deemed important for the selection of habitat types and features by elk. Therefore we believe reasonable variables were investigated and should provide Table 4. Mean attributes at elk feeding (n=96) and control plots (n=96) in west-central Alberta during the winter of 1994 — 1995 Elk Feeding Sites Control plots M+ SE Mean + SE P Value* Non-spatial Variables Canopy Closure (%) 31.34 + 3.45 63:12: 3-355 < 0.001 Grass Cover (%) 59.73 + 3,37 32.61 + 3.20 < 0.001 Herb Cover (%) 19.00 + 1.16 19.85 + 1.01 0.574 Shrub Cover (%) ia.Al +4250 26.25 + 1.95 < 0.001 Spruce, Pine, & Fir in the canopy (%) 28.17 + 4.20 44.70 + 4.06 0.004 Stem Density (#/ ha < 1000) foi 2019 3.24 + 0.29 < 0.001 Tree Height (m) 5.46 + 0.72 12.44 + 0.78 < 0.001 Spatial Variables Major Road 1093 + 66.06 1088 + 64.50 0.829 Minor Road 601 + 70.05 635 + 72.24 0.139 Unimproved Access 102 + 11.30 164 + 12.34 < 0.001 Thermal Cover 160 + 14.35 158 + 18.25 0.932 Hiding Cover 71 + 7.68 48 + 8.06 0.024 * P-values are the result from paired t-test. 188 THE CANADIAN FIELD-NATURALIST Vol. 116 Table 5. Mean attributes at elk bedding (n=62) and control plots (n=62) in west-central Alberta during the winter of 1994 — 1995 Elk Feeding Sites Control plots M + SE Mean + SE P Value? Non-spatial Variables Canopy Closure (%) 64.68 + 4.03 71.74 + 4.01 0.157 Grass Cover (%) 34.95 + 3.30 DI 3IDEZ 2 0.016 Herb Cover (%) 16.85 + 0.89 Lh, 7A: 29 0.56 Shrub Cover (%) 2ST E76 28.20: 213 0.698 Spruce, Pine, & Fir in the canopy (%) 38.53 + 5.07 54:21 25215 0.023 Stem Density(#/ha X 1000) 2.0352 017 3172027 0.001 Tree Height (m) 13.38 + 0.87 13.75 20:78 0.701 Spatial Variables Major Road 1244 + 87.46 1229 + 84.34 0.550 Minor Road 763 + 69.90 ISL Is 0.830 Unimproved Access 165 + 19:59 155 = 1950 0.684 Forage Stands 71+11.14 Ot 15:63 0.880 * P-values are results from paired t-test. meaningful insight into the habitat selection patterns of elk at multiple scales. During winter, vegetation is at its lowest quality and quantity and therefore elk should select areas containing the highest quantity of available forage. Selecting areas of high forage concentration would reduce energy expenditures associated with locomo- tion through snow and decreased temperatures (Geist 1982). Previous studies have shown elk prefer to graze on grasses (site selection) when snow depths do not impede foraging (Nelson and Leege 1982; Morgantini 1988). The selection of grass/meadows as foraging sites at the stand level has been well doc- umented (Craighead et al. 1973; Morgantini and Hudson 1979; Irwin and Peek 1983; Jenkins and Wright 1986; Morgantini 1988). The proportion of a home range as forage or habitat diversity (e.g., patch size and density) may be variables that are represen- tative of a landscape scale for elk. Previous studies have indicated habitat selection by elk at the land- scape level. Irwin and Peek (1983) concluded that forage density strongly influenced the size and loca- tion of elk home ranges in Idaho. In dry shrub steppe environments, larger home ranges may compensate for low forage availability (McCorquodale et al. 1989). Black et al. (1976) proposed that optimal elk habitat consists of 40% of the land base as cover and 60% as food in the proper size and spatial arrange- ment. As an ecotone species, elk select heteroge- neous areas for winter home ranges providing the opportunity to feed in grass meadows while taking advantage of the adjacent areas for cover (Geist 1982). The present study confirms previous reports on selection of forage characteristics by elk at different scales. At the site scale elk selected open areas with high grass cover for feeding when compared to bed- ding locations and available habitat. At the stand level elk selected grass/meadows due to their con- centration of grasses while utilizing all other habitat types in proportion to their availability. The results of the landscape level analysis are less conclusive. If habitat diversity, which is represented by patch size and density, is representative of a need for a hetero- geneous environment to meet different life requi- sites, then elk did exhibit landscape level selection. Though not significant in comparison to available home ranges, elk home ranges showed little variation in the proportion classified as forage even though there was a wide range of home range sizes. This suggests that forage maybe an important factor in determining home range size and location in our study. Closed canopies can reduce thermoregulatory costs, reduce snow depths, and provide security for elk (Peek et al. 1982; Skovlin 1982; Peek and Scott 1985). Thermal cover is provided by stands that pro- tect against extreme temperatures, whereas hiding cover provides a sense of security from predators and human disturbance. Much debate has centered around whether elk select closed stands for their thermal or hiding cover value (Geist 1982; Cook et al. 1998). Beall (1976) concluded that elk selected closed stands to reduce heat loss, whereas Peek and Scott (1985) believed that thermal cover would have an insignificant effect on energy requirements. They concluded that security cover was important in the presence of human disturbance (Peek and Scott 1985). In western Alberta, Morgantini and Hudson (1979) concluded that habitat selection by elk in winter was not a response to the thermal environ- ment, but governed by human disturbance. Our results suggest that elk did not select for ther- mal cover requirements at any scale. At the land- scape level the proportion of the home range classi- fied as thermal cover was comparable to that of the 2002 AHR, while at the stand level those habitat types that would represent thermal cover (e.g., closed mature or closed old-growth) were used in proportion to their availability. At the site scale elk selected bed- ding sites whose structural characteristics would be considered of lower thermal cover quality than the available habitat. The need for hiding cover by elk is well docu- mented (Hershey and Leege 1976; Morgantini and Hudson 1979; Peek and Scott 1985). Most studies have described the need for hiding cover in regards to human disturbance and, in particular, roads. The avoidance of habitat near open roads by elk has been extensively documented (Hershey and Leege 1976; Lyon 1979; Rost and Bailey 1979). Most studies have established avoidances of roads in terms of a distance, but Lyon (1983) determined avoidance or loss of habitat in terms of road density. Habitat effectiveness decreases dramatically as the density of roads increases such that an area with a road density greater than 0.62 km/km/? is reduced by 40% (Lyon 1983). The results of selection by elk at multiple scales in terms of security requirements are less conclusive than those found for forage and thermal cover requirements. At the landscape level there was not a significant difference in the proportion of hiding cover comprising either the elk home ranges or AHR. This is not surprising as the study occurred in an area that is predominantly forested. We would argue that selection at the landscape level did occur in terms of hiding cover requirements based on the results of the road density analysis. Road densities for the elk home ranges were significantly less than those of the AHR. In addition the densities are below the 0.62 km/km/? density established by Lyon (1983). Previous studies of habitat selection have been conducted at a single scale, with results being extrapolated to higher or lower levels (Pedlar et al. 1997). The appropriateness of extrapolating results to other levels has not been examined. Our results involving selection by elk of forage characteristics at the three levels would support the extrapolation of results to higher or lower levels. Based on our analy- sis, conclusive results for habitat selection were observed at the site and stand scale with inconclusive results at the landscape scale. This suggests that habitat selection, in terms of forage requirements, are bottom up as opposed to top down. There was no selection detected at any level by elk to meet the needs of the life requisite of thermal cover, therefore extrapolation to other levels would be appropriate (though unnecessary). The appropriateness of extrap- olating the results of the hiding cover analysis is less conclusive. Selection of habitat characteristics to meet the requirement of hiding cover was not evi- dent at the stand or site level, but may have been masked by the selection at the landscape level. JONES AND HUDSON: WINTER HABITAT SELECTION BY AMERICAN ELK 189 During different times of the year and under increased levels of human disturbance selection for hiding cover may be more evident at the stand and site level. We acknowledge that our conclusions are based on the analysis of habitat selection data from one winter with lower than average precipitation levels. Because of this we did not examine the influence that snow depth had on habitat selection. It was also beyond the scope of our study to examine other factors (e.g., predation, inter and intra-specific com- petition) that can influence habitat selection. To over- come these limitations, further investigation is war- ranted to determine how these other factors change habitat selection patterns and our conclusions. The value of our study is to show resource managers the need to examine habitat selection patterns across more than one level to better manage elk. Acknowledgments We gratefully acknowledge the financial support of the Foothills Model Forest, without which this project could never have been completed. In addition we would like to thank the Hinton Fish and Game Association and the Rocky Mountain Elk Foundation for their contributions. Equipment and logistic sup- port was provided by Weldwood of Canada Inc. (Hinton Division), Alberta Environmental Protec- tion, Fish and Wildlife Division (Edson), NOVA Gas Corporation, and The Environmental Training Centre. We thank F. Nelson of the Athabasca Ranch for permission to travel across her property. To D. Jewison, N. Gataiant, S. Kathnelson, W. Jones, and the many volunteers who helped in the collaring of elk and the collection of field data. Special thanks to M. Todd and D. Farr for their timely advice and guidance. 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Pages 104-127 in Wildlife habitats in managed forests; the Blue Mountains JONES AND HUDSON: WINTER HABITAT SELECTION BY AMERICAN ELK 191 of Oregon and Washington. Edited by J. W Thomas. U. S. D. A. Forest Service Handbook Number 553, Washing- ton, D.C. Thompson, M. J., R. E. Henderson, T. O. Lemke, and B. A. Sterling. 1989. Evaluation of a collapsible clover trap for elk. Wildlife Society Bulletin 17: 287-290. Trimble Navigation. 1992a. GeoExplorer operation man- ual. Trimble Navigation, Sunnyvale, California, USA. Trimble Navigation. 1992b. Pathfinder system general ref- erence. Trimble Navigation, Sunnyvale, California, USA. White, G. C., and R. A. Garrott. 1990. Analysis of wildlife radio-tracking data. Academic Press, San Diego, California, USA. Received 31 March 2001 Accepted 20 May 2002 Notable Vascular Plants from Alaska in Wrangell-St. Elias National Park and Preserve with Comments on the Floristics Mary B. Cook! and CARL A. ROLAND? \Wrangell-St. Elias National Park and Preserve, P.O. Box 439 Copper Center, Alaska 99573, USA 2Denali National Park and Preserve, P.O. Box 9, Denali Park, Alaska 99755, USA Cook, Mary B., and Carl A. Roland. 2002. Notable vascular palnts from Alaska in Wrangell-St. Elias National Park and Preserve with comments on the floristics. Canadian Field-Naturalist 116(2): 192-304. An inventory of the vascular flora north of the Bagley Icefield in Wrangell-St. Elias National Park and Preserve, Alaska, U.S.A., was conducted from 1994 to 1997. The objectives of the inventory were to assess the genetic diversity of the region, identify rare taxa and areas of phytogeographic interest and to assist park managers with planning and environmen- tal compliance. The purpose of this paper is to present annotations for 212 of the most notable taxa with their Alaska- Yukon and Park distribution maps and to summarize the floristics of the Park. 832 species (887 taxa) were documented as occurring within the Park (approximately 54% of the Alaskan flora). Significant results from the inventory include: (1) nine additions to the flora of Alaska (Arabis calderi G. A. Mulligan, A. codyi G. A. Mulligan, A. drepanoloba Greene, A. lemmonii S. Wats., Carex petasata Dewey, Draba lonchocarpa Rydb. var. thompsonii (C. L. Hitchc.) Rollins, Festuca minutiflora Rydb., Najas flexilis (Willdenow) Rostkov. & Schmidt and Trichophorum pumilum (Vahl.) Schinz & Thell. var. rollandii (Fern.) Hultén); (2) 331 range extensions; (3) the addition of 40 Alaska Natural Heritage plant taxa with a state rank of three or less for a total of 69 rare plant taxa in the Park’s flora, and (4) the addition of 214 species to the Park’s flora. The composition of the flora in percentages, with the rare flora in parenthesis, is: 25.8 (8.3) circumpolar, 10.8 (9.7) incompletely circumpolar, 32.2 (38.9) North American, 23.2 (11.1) amphiberingian, 0.9 (1.4) amphiatlantic and 7.1 (26.4) Alaska-Yukon endemic. The Alaska-Yukon and amphiberingian endemic elements are dominant in the Alaska Range, the Wrangell Mountains and in the Tanana, White and Copper River basins whereas the Cordilleran endemic element is dominant in the St. Elias and Chugach Mountains and Chitina River basin. These trends correspond to our understanding of plant migration after the Pleistocene Epoch from refugia in the Upper Yukon Valley, the Alaska Range and Beringia, the northern part of the Park being closest to these migration corridors, whereas the St. Elias and Chugach Mountains are at the northern end of the North American cordilleran migration corridor. Endemics may also have evolved from relict populations in unglaciated areas within the Late Wisconsin ice sheet adjacent to Lake Ahtna, on coastal refugia and on exposed sites in the dry northern interior of the Park bordering the Tanana Valley and the southeastern edge of Beringia. The rare flora was distributed somewhat evenly through the mountain ranges of the Park, but unevenly through- out the basin regions of the Park where the Chitina River basin had two to four times the number of rare species found in the other basins. There was a trend for rare plants to occur in the alpine zone, above 1200 m elevation, in xeric sites, in the alpine-herb talus slope plant community, on southerly aspects and on slopes of 20-40 degrees. Although the Park is protected, infrastructure development, visitor use in the backcountry in communities most likely to harbor rare plants, and global changes to the ecosystems are increasing. This inventory enhanced our understanding of the genetic diversity, phytogeography and distribution of our flora, however, there are still large areas in the Park that have not been surveyed, particularly in the coastal region, and we lack sufficient knowledge about most rare species (distribution, life history and population ecology) to develop protection plans. Key Words: Vascular plants, flora, rare plants, floristics, biogeography, range extensions, National Park, Alaska. Wrangell-St. Elias National Park and Preserve encompasses a unique cross section of boreal and coastal ecosystems in south-central Alaska with floristic influences from Beringia, the Yukon, the arctic and the Pacific Mountain systems (Figure 1). There is a high diversity of plant communities in this region due in part to the large size of the Park (5.3 million hectares), the three climatic zones it covers (interior, montane and transitional) and the wide variety of landforms and lithologies found within its boundaries. Eric Hultén, in his history of botanical exploration in Alaska, identified the Wrangell-St. Elias region as one of three areas in Alaska that was poorly known (Hulten 1940). The distribution maps in his flora of Alaska (Hultén 1968) show a void in the Wrangell-St. Elias region for many boreal-montane and arctic- alpine taxa that would be expected to occur here. Published floristic studies within the Park were limit- ed to a few geographic areas: Skolai Creek, Chitistone Pass and the Chitistone River (Murray 1968, 1971); Russell, Sheep and Guerin Glaciers (Murray 1971); Chitistone, Skolai and Frederika Valleys (Scott 1968, 1974); Long Glacier and Dadina River (Saltmarch 1978) and Bonanza Ridge near Kennicott (Nordell and Schmitt 1977). Other collections were made prior to the establishment of the Park, but these were pri- marily along the two roads into the Park. Collections made from 1984 to 1992 during Park vegetation stud- ies documented 121 range extensions, twenty-two new rare plant species and verified that large areas of the Park were still unsurveyed. 192 2002 ASSL =) S 145° 30' W COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 193 62° 30'N MALASPINA GLACIER wees Park Boundary FIGURE 1. Map of study area showing major geographical features of Wrangell-St. Elias National Park and Preserve, Alaska. The Park was established in 1980 under the Alaska National Interest Lands Conservation Act (ANILCA, Public Law 96-487). An inventory of the plants occurring within the Park is necessary to ful- fill the following management objectives of the Park as stated in the Final Environmental Statement (Alaska Planning Group 1973): 1. Ensure retention of the magnificent Wrangell- St. Elias landscapes and living systems in a natural state. 2. To the extent possible, allow the natural fluctu- ations and equilibrium of self-regulating ecosystems to continue unimpeded. 3. Develop and implement a viable research program to provide basic information required for effective Park management. 4. Lay early emphasis on identification of espe- cially fragile areas through appropriate research. The Natural Resources Management Guideline for the National Park Service, NPS-77 (USDI 1991) states that, “A basic vegetation inventory is the first step in vegetation management”, and the General Management Plan for the Park mandates that the presence and extent of endangered species of flora and fauna be determined (USDI 1986). Vegetation in the Park is subject to alteration from recreational and subsistence uses including mining, timber harvest, grazing, all-terrain vehicles, hiking, climbing and camping. Backcountry use is increasing steadily as the Park infrastructure is developed and as the public seeks to experience new premier wilderness areas. Identification of sensitive taxa, their habitats, and unique floristic associations was deemed necessary by the National Park Service for the preparation of backcountry management plans, environmental assessment and compliance and for the monitoring of subsistence uses. We conducted an inventory of the vascular flora of selected areas north of the Bagley Icefield within the Park from 1994 to 1997. The objectives of the 194 inventory were to: increase our understanding of the ecological history, genetic diversity and biogeogra- phy of the Park’s flora; identify populations of rare taxa, unique floristic associations and areas of phytogeographic interest in order to protect the biodiversity of the natural systems within the Park, and to provide the structure for an ongoing assess- ment of the Park’s flora. The purpose of this paper is to present the collection data for the most notable range extensions along with Park and Alaska- Yukon distribution maps and to summarize the floristics of the Park. Study Area Four large mountainous regions define the land- scape of the Park: the Wrangell Mountains, the St. Elias Mountains, the Chugach Mountains and the Alaska Range. The Wrangell Mountains, a volcanic range with peaks reaching 4995 m, dominate the landscape in the center of the Park. The most com- mon bedrock in the Wrangell Mountains is Wrangell Lava, although the southern Wrangells have large exposures of Chitistone and Nizina limestone, Nikolai Greenstone and metamorphosed sedimentary rocks (MacKevett 1978; Richter 1976). The St. Elias Mountains straddle the border with the Yukon Territory and are even more precipitous and heavily glaciated than the Wrangell Mountains with peaks rising to 5489 m. The most common surface rock types in the St. Elias Mountains are Wrangell lava, Nikolai greenstone and marine sedimentary rocks of the Hasen Creek formation such as argillite, chert, conglomerate, limestone and shale (MacKevett 1978). The Chugach Mountains are heavily glaciated due to their proximity to the Gulf of Alaska and extensive glaciation during the Pleistocene as indi- cated by the steep-sided U-shaped valleys, obvious trimlines, hanging glaciers and sparsely vegetated morainal features. The lithology of the Chugach Mountains is dominated by rocks of the Valdez group, a marine sedimentary unit composed primari- ly of graywacke, argillite, slate and phyllite (MacKevett 1978). North of the Wrangell Mountains lies the eastern arc of the Alaska Range which lies south of the Denali Fault and is represented within the Park by the Mentasta and Nutzotin Mountains. The lithology of the eastern Alaska Range is largely composed of marine sedimentary rocks with expo- sures of Nikolai Greenstone and thin-bedded limestone (Richter 1976). The Copper, Chitina, White and Tanana (includ- ing the Nabesna and Chisana Rivers) are major inte- rior river drainage systems found within the Park. These drainage basins are mantled with quaternary surficial deposits of diverse origins. Surficial deposits include drift from the Wisconsin and older glaciations, Eolian deposits, Holocene alluvium and lacustrine sediments. The principal depositional fea- ture in the Copper River basin was the presence of THE CANADIAN FIELD-NATURALIST Vol. 116 the huge pro-glacial Lake Ahtna, which formed behind an ice dam at the confluence of the Copper and Chitina Rivers during the Pleistocene Epoch. Lake Ahtna occupied more than 520 000 hectares of the Copper River basin to a maximum elevation of 800 m on the flanks of the Wrangell Mountains (Ferrians 1989). In contrast to the large expanses of open, low elevation terrain present in the Copper River basin, the upper Nabesna and Chisana Rivers occupy relatively narrow valleys. The valley floors of these braided rivers lie within the active flood- plain of the streams. The surficial deposits in the large, open valleys of the Chitina and White Rivers are a mix of alluvium on the floodplains and glacial outwash and drift from the Wisconsin glaciation on surfaces that have not been reworked by fluvial processes. Large areas in the White River valley are blanketed with volcanic ash from the eruptions of Mt. Churchill that occurred 1900 and 1250 years ago. Precipitation in the Park ranges from a yearly average of 338 cm at Yakutat, located in the mar- itime climatic zone, to 20 cm at Slana in the interior climatic zone. Temperatures on the coast are mild ranging from a mean daily high of 15°C to a mean daily low of —9°C whereas temperatures at Slana range from a mean daily high of 20°C to a mean daily low of —25°C. Five ecoregions and three transitional areas are found within the Park (Gallant et. al. 1995). These are: (1) the coastal Western Hemlock-Sitka Spruce forest ecoregion which is found on the Malaspina Forelands between Icy and Yakutat Bays; (2) the Pacific Coastal mountain ecoregion which includes the Chugach Mountains, southern St.-Elias Mountains and the Bagley Icefield; (3) the Wrangell Mountain ecoregion, demarcated by the Wrangellia Terrane; (4) the Copper Plateau ecoregion, the area of pro-glacial Lake Ahtna; (5) the Interior Highlands ecoregion, delineated by the Yukon-Tanana Terrane in the northeast corner of the Park; (6) the Alaska Range transition ecoregion encompassing the Nutzotin and Mentasta Mountains; (7) the interior bottomlands transition ecoregion, the region north of the Nutzotin Mountains in the Tanana Lowlands, and (8) the Duke Depression transitional ecoregion, the area encompassing the White River along the Duke River fault. Most of the Park was covered by the Cordilleran ice sheet or by the large pro-glacial Lake Ahtna during the peak of the Wisconsin glaciation (Clague 1992; Hamilton 1994; Hamilton and Thorson 1983). The major river valleys became ice-free and Lake Ahtna drained with the retreat of the glacier occupy- ing the mouth of the Copper River canyon by 9.4 ka (Denton 1974; Hamilton 1994). There may have been unglaciated areas within the Late Wisconsin ice sheet adjacent to Lake Ahtna (Hamilton and Thorson 1983), on coastal refugia and on exposed sites in the 2002 dry northern interior of the Park bordering the Tanana Valley and the southeastern edge of Beringia (Hopkins 1967). Previous Botanical Collections Eight significant plant collections were made within the Park prior to its establishment in 1980. William L. Poto collected 175 specimens primarily along the Mt. Drum Trail in the central Wrangell Mountains in 1902 as a member of the United States Geological Survey (USGS) Mt. Wrangell and Central Copper River Region Exploration Expe- dition (Poto 1902*; Mendenhall and Schrader 1903; Mendenhall 1905; Hultén 1940). Frank Charles Schrader and G. H. Hartman collected in the north- ern Wrangell Mountains between the Nabesna and Copper Rivers the same year as Poto as part of this expedition. The specimens of Poto, Schrader and Hartman are at the U.S. National Herbarium. David W. Eaton, DeLorme D. Cairnes and H. F. Lambart collected along the Yukon border, in the Park in the Wrangell and St. Elias Mountains from 1909-1913 as part of the Yukon-Alaska International Boundary Survey (Cairnes 1911, 1914; Macoun 1914; Hultén 1941-1950; Green 1982). The collections of Eaton are at the U.S. National Herbarium, those of Cairnes and Lambart are at Canadian Museum of Nature in Ottawa. Hamilton M. Laing, a biologist with the Department of Mines of British Columbia who was a member of the Mt. Logan expedition in 1925, collected 243 specimens at the head of the Chitina River primarily in the St. Elias and Chugach Ranges. His collection was classified by A. E. Porsild and is housed at the Canadian Museum of Nature in Ottawa (Porsild 1939; Lambart 1926a, 1926b; Hultén 1941-1950). David F. and Barbara Murray collected at May Creek, Nizina, Chitistone Pass, Skolai Pass, Guerin Glacier, Russell Glacier and Sheep Glacier in the Wrangell-St. Elias Mountains from 1966 to 1981 (Murray 1968, 1971). Their collections are at the University of Alaska Fairbanks Museum and at the Canadian Museum of Nature in Ottawa. Richard W. Scott collected at Frederika Glacier, Chitistone Pass and Snag Glacier in the Wrangell Mountains in 1967 and 1968 while conducting an ecological phytogeo- graphical study of the southeastern Wrangell Mountain flora (Scott 1968, 1974). Scott’s collection is at the University of Alaska Fairbanks Museum and at the University of Michigan. Olle Nordell and Alf Schmitt collected 207 specimens at Kennicott and Bonanza Ridge in the Wrangell Mountains in 1976 (Nordell and Schmitt 1977). Their specimens are at Lund and the University of Alaska Fairbanks Museum. Ransom Saltmarch collected 76 specimens in 1978 on the slopes of Mt. Wrangell while con- *See Documents Cited section. COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 195 ducting research for a novel (Saltmarch 1978). His specimens were reviewed by D. F. Murray (ALA). An additional 36 individuals collected primarily along the two roads into the Park or on the roads adjacent to the Park. Among these collectors are: Edwin F. Glenn (1899), Martin Woodlock Gorman (1898 and 1899), E. L. Blaschke (1820), Arthur James Collier (1902), Adolphus Washington (1905), I. E. Diehl (1908), Walter Harrison Evans (1909), Bayne-Beauchamp Expedition (1932), William Albert Setchell (1932), Frits Warmolt Went (1934), Jacob Peter Anderson (1935 and 1944), Elisabeth Kol (1936), H. M. Raup (1944), Artheme-Antoine Dutilly and Ernest LePage (1945-1947), Eric Hultén (1961), M. Sharrock (1962), George Argus (1967), R. Pegau (1968, 1970), and Leslie Viereck (1957-1980). The only collections made along the Park’s coast were eight made by Frederick Funston in 1892 near Esker Stream and Manby Point on the Malaspina Forelands, five made by the Harriman Expedition at the Hubbard Glacier in 1899 and numerous Salix specimens collected by George Argus near Esker Stream (Coville and Funston 1896; Hultén 1941-1950; Argus 1967). Park staff collected 1145 specimens at 215 unique localities throughout the Park from 1982 to 1994 during research and resource management studies. Methods We conducted a reconnaissance inventory north of the Bagley Icefield from 1994-1997. Site selection focused on surveying a range of plant communities, lithologies (such as calcareous, ultramafic, basalt, ash and greenstone) and landforms (such as south- facing bluffs, sand dunes, warm springs and flood- plains). We also focused site selection on azonal communities (scree slopes, wetlands and aquatics) and those areas where we predicted rare and endemic species would occur. In all, 239 sites were surveyed and approximately 7000 specimens were collected. Specimens were critically examined by the authors, Alan Battan, David F. Murray and Carolyn Parker at the University of Alaska Museum and the following specialists: George W. Argus (CAN) — Salix; Reidar Elven (O) — Draba lactea and Cerastium regelii; Mark Egger (WTU) — Castilleja; Barbara Ertter (UCB) — Potentilla diversifolia and P. drummondii; Donald Farrar (ISC) — Botrychium; Signe Frederickson (C) — Festuca; G. A. Mulligan (DAO) — Arabis and Draba; David F. Murray Papaver and Carex section Atratae; and Marcia Waterway (MTMG) — Carex laxa. We developed a set of relational databases in which the biogeography, taxonomy and rarity of each species was recorded and site characteristics for each rare plant collection was documented and evaluated. Distribution maps were prepared for selected taxa by assembling the following locality 196 data into a Geographic Information System: all collections from this inventory and previous Park collections, specimen records from the University of Alaska Fairbanks Museum Plant Documentation Center and published stations from regional floras and monographs (Aiken and Darbyshire 1990; Argus 1973; Argus 2000; Bayer 1993; Cody 1996; Cody et. al. 1998; Hultén 1941-1950; Hultén 1968; Porsild and Cody 1980). The distribution maps are at two scales, one encompassing all of Alaska and the Yukon Territory including adjacent parts of Canada and one focused on the Park. Nomenclature follows Cody (1996) for scientific names and Kartesz and Meacham (1999*) for common names. Results and Discussion Taxonomic Composition There are 832 species documented by vouchers within the Park, with a total of 887 taxa including intraspecific taxa. The flora of the Yukon Territory has approximately 1199 species, the flora of Alaska has approximately 1535 species and the two regions combined have approximately 1654 species with 1202 species in common (Kartesz and Meacham 1999). The number of species in the Park’s flora is therefore approximately 54% of the Alaskan flora and 69% of the Yukon Territory flora. For compari- son with other protected areas, Denali National Park and Preserve in Alaska reported 684 taxa in 2000 and Kluane National Park in the Yukon Territory has recorded 704 taxa (656 species) (Caswell 2002*). The flora of the Park is distributed among 81 families and 258 genera. Eight of the species are not native to Alaska. The most important plant families within the flora are the Cyperaceae with 100 species (104 taxa), the Poaceae with 69 species (84 taxa), the Asteraceae with 68 species (75 taxa) and the Brassicaceae with 66 species (71 taxa). Phytogeography The Wrangell-St. Elias region is located within the distribution patterns of the major North American floristic elements, hence the plant species here have diverse origins and histories. They include species endemic to the area, broadly distributed boreal forest plants, arctic-alpine species, endemics of the western mountains and amphiberingian taxa of Asian origin. We recognize the following floristic elements within the vascular flora of the Park: I. Circumpolar species — 25.8% of the flora (226 species). This group includes those broadly distributed species that occur on all circumpolar continents including both Asiatic and European regions of Eurasia, Greenland and North . America. We further divide this group of plants into boreal species primarily found in lowland and montane habitats and arctic/alpine taxa that generally occur in treeless landscapes in the arctic or alpine regions. A third element of the circum- polar flora is a group of more wide-ranging species with broad ecological amplitudes that occur across both arctic and THE CANADIAN FIELD-NATURALIST* Vol. 116 boreal zones. The ratio of these elements within the circum- polar flora and examples of taxa from each category are: (1) Arctic-alpine species (10.6%) — Cardamine bellidi- folia, Carex lachenallii, Draba fladnizensis, Erigeron humilis, Saxifraga oppositifolia and Silene acaulis. (2) Boreal-montane species (6.7%) — Carex tenuifolia, Linnaea borealis and Rosa acicularis. (3) Widespread species (8.5%) — Carex capillaris, Empetrum nigrum, Equisetum arvense and Poa glauca. II. Incompletely circumpolar species — 10.8% of the flora (94 species). Species in this group have distributions that are very similar to the circumpolar plants of boreal distri- bution, but are not known from either Greenland or Europe. Examples of incompletely circumpolar species include Calypso bulbosa, Carex limosa, Chamaedaphne calycula- ta, Equisetum fluviatile, and Moehringia lateriflora. III. North American species — 32.2% of the flora (282 species). Taxa in this group have distributions that are gen- erally restricted to North America. Two additional groups of species with more narrow distributions can be identified within the North American element: cordilleran species, which occur in the western mountains, and Pacific coastal species, which are generally restricted to the coast ranges and the west coast of North America. (1) Arctic-alpine species (2.1%) — Erigeron composi- tus and Primula egaliksensis (2) Boreal-montane species (11.4%) — Shepherdia canadensis and Viburnum edule (3) Cordilleran species (10.2%) — Luetkea pectinata and Salix commutata (4) Pacific coastal species (6.2%) — Lupinus nootka- tensis and Epilobium luteum (5) Widespread species (2.4%) — Solidago multiradiata IV. Amphiberingian species — 23.2% of the flora (203 species). These are species that occur in North America and northern Asia but are not known from either Greenland or Europe; hence their center of distribution generally lies within Beringia. We separate the species within the amphiberingian biogeographic element into the following categories: (1) Arctic-alpine species (8.9%) — Artemisia arctica and Senecio atropurpureus (2) Boreal-montane species (4.3%) — Achillea sibirica and Boschniakia rossica (3) Pacific coastal species (5.0%) — Carex macro- cheata and Fritillaria camschatcensis (4) Widespread species (5.0%) — Parrya nudicaulis and Carex podocarpa V. Amphiatlantic species — 0.9% of the flora (8 species). Taxa in this group of species have distributions that include North America, Greenland and Europe but are not known from Asia. Carex nardina and Draba crassifolia are exam- ples of this element. VI. Alaska-Yukon endemic species — 7.1% of the flora (62 species). These are species that are restricted to Alaska and Yukon Territory, and sometimes extend into neighbor- ing regions of Northwest Territiories and British Columbia. (1) Arctic-alpine species (2.3%) — Thlaspi arcticum and Smelowskia borealis (2) Boreal-montane species (2.2%) — Salix setchel- liana and Artemisia alaskana (3) Cordilleran species (1.4%) — Synthyris borealis and Stellaria alaskana 2002 (4) Pacific coastal species (1.3%) — Castilleja una- laschensis and Salix stolonifera The composition of our flora is very similar to that described by George Douglas for the southwest Yukon (Douglas 1974). The northerly elements (circumpolar, incompletely circumpolar, North American arctic-alpine, amphiatlantic and amphi- beringian) comprise 64% of both floras whereas the southern elements (Cordilleran and Pacific Coast) contribute 16% to each flora. The North American boreal-montane group contributes 14% to the south- west Yukon flora and 11% to the Wrangell-St. Elias flora. The southwest Yukon flora had eight non- native species in 1974 as does the current Wrangell- St. Elias flora. Alaska- Yukon endemics comprise 7% of our flora and 5% (32 species) of the southwest Yukon flora reported by Douglas (1974). Lausi and Nimis (1985) documented 23 Alaska-Yukon endemics for the southern Yukon in a phytosociolog- ical study along the Alaska Highway. Porsild (1951) estimated that amphiberingian species comprise approximately 33% of the Alaska-Yukon flora. Douglas noted the southward decrease in this element for the southwest Yukon flora (17%). The slight increase in the contribution of this element to the Wrangell-St. Elias flora (23%) would support this gradient. We found the ratio of arctic and coastal elements of the southern Wrangell Mountain flora to be similar to that described by R. W. Scott in his phyto- geographical study of the southeast Wrangell Mountains (Scott 1974). Scott’s analysis described a total of 291 species with 21% arctic, 7% coastal, 55% montane and 17% boreal. Our analysis resulted in 334 species for the southern Wrangells with 39% arctic- alpine, 23% boreal-montane, 12% cordilleran, 6% Pacific coastal, 20% widespread and 0.3% introduced. OAlaska-Yukon Amphiberingean COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 197 Distribution of the Endemic Elements within the Park We evaluated the ratio of the endemic elements (Alaska- Yukon, amphiberingian, cordilleran and Pacific coastal) within each of the mountain and basin regions in the Park to determine if the composi- tional patterns are related to the ecological history of the region (Figures 2 and 3). The Alaska- Yukon and amphiberingian endemic floras are dominant in the Alaska Range (39% and 40% respectively), the Wrangell Mountains (28% and 42%) and in the Tanana (58% and 18%), White (39% and 52%) and Copper River basins (43% and 35%). This trend cor- responds to our understanding of plant migration after the Pleistocene Epoch from refugia in the Upper Yukon Valley, the Alaska Range and Beringia, the northern part of the Park being closest to these migra- tion corridors (Hultén 1937; Murray, et. al. 1983; Yurtsev 1963; Murray 1995). The Cordilleran endemic flora is dominant in the St. Elias Mountains (45%), Chugach Mountains (37%) and in the Chitina River basin (38%) as would be expected since the St. Elias and Chugach Mountains are at the northern end of the North American cordilleran. Several of our rarest species are cordilleran endemics reaching their northern extent in the Park (Arabis calderi G. A. Mulligan, A. Jemmonii S. Wats, Carex petasata Dewey, and Festuca minutiflora Rydb). The Pacific coastal endemic element decreases notably from south to north in both the mountain and basin ranges. This trend may indiciate that migration north from the coast is impeded by the mountain ranges or by the availability of suitable habitat. The strong latitudinal gradient may also indicate that the Pacific coastal species represent a younger element in our flora when compared to the other three elements, which are more evenly distributed throughout the basin and Alaska Range (n=130) Wrangells (n=207) St. Elias (n=53) Chugach (n=152) Mountain Regions FIGURE 2. Ratio of endemic taxa within mountain regions of Wrangell-St. Elias National Park and Preserve, Alaska. 198 mountain ranges. Endemics may also have evolved from relict populations in unglaciated areas within the Late Wisconsin ice sheet adjacent to Lake Ahtna (Hamilton and Thorson 1983), on coastal refugia and on exposed sites in the dry northern interior of the Park bordering the Tanana Valley and the southeast- ern edge of Beringia (Hopkins 1967). The Rare Flora Sixty-nine species in the Park’s flora have an Alaska Natural Heritage state rank less than three and are treated as rare species by the National Park Service, 40 of these are new to the Park’s flora. None of the rare species are considered threatened or endangered by the U.S. Fish and Wildlife Service. However, three species were listed as Species of Concern by the U.S. Fish and Wildlife Service in 1995 (Cryptantha shackletteana L.C. Higgins, Carex laxa Wahlenb. and Taraxacum carneocoloratum Nels.). The most important families in the rare flora are the Brassicaceae with 17 species, the Cyperaceae with 13 species and the Caryophyllaceae with seven species. The three dominant classes for the distribu- tion of our rare flora by form of rarity (Rabinowitz 1981) are: constantly sparse and geographically restricted in a specific habitat (30 species), locally abundant in a specific habitat but restricted geo- graphically (17 species) or constantly sparse in a specific habitat but occurring over a large range (12 species). Trends in the distribution of the rare flora by site characteristics are summarized by the following values. The number in parenthesis is the percent of total sites with that particular value for a site charac- teristic evaluated using 239 sites and 423 rare plant collections. 82% (37%) of all rare plant collections were made in the alpine zone; 78.5% (41.8%) were above 1200 m; 61% (37%) were in a xeric moisture class, and 57% (19%) were on alpine herb-talus slopes. 57% of all plant collections had a slope of 20-40 degrees and 45% were collected on southerly aspects. The remaining aspect classes were distribut- ed approximately equally (north — 17%, east — 19% and west — 19%). The rare flora appears to be distributed relatively evenly throughout the mountain ranges (> 1067 m) in the Park. The Alaska Range has 32 rare species, six of these being unique to the Alaska Range; the Wrangell Mountains have 33 species with three species unique to the Wrangell Mountains; the St. Elias Range has 19 rare species, two of which are unique to the St. Elias Range; and the Chugach Mountains (including the Granite Range) have 28 rare species, eight of which are unique to the Chugach Mountains. The distribution of rare plants is uneven for the river basins in the Park. The Chitina River Basin has 32 rare species, six of which are unique to this basin. The Tanana River has 13 THE CANADIAN FIELD-NATURALIST- Vol. 116 rare species, two being unique; the Copper River basin has 13 rare species, two being unique, and the White River basin has eight rare species, with no species unique to that basin. The biogeographic composition of the rare flora with examples of species from each element follow. I. North American species — 38.9% (28 species). (1) Arctic-alpine (1 species) — Primula egaliksensis (2) Boreal-montane (8 species) — Agoseris glauca, Carex parryana, Eriophorum viridi-carinatum and Maianthemum stellatum (3) Cordilleran (19 species) — Arabis calderi, A. lem- monii, Carex petasata and Festuca minutiflora If. Alaska-Yukon endemics — 26.4% (19 species). (1) Arctic-alpine (6 species) — Thlaspi arcticum and Douglasia arctica (2) Boreal-montane (3 species) — Cryptantha shacklet- teana, Lupinus kuschei and Salix setchelliana (3) Cordilleran (6 species) — Arabis codyi and Taraxa- cum carneocoloratum If. Amphiberingean species — 11.1% (8 species) (1) Arctic-alpine (5 species) — Festuca lenensis, Stel- laria umbellata and Trisetum sibiricum (2) Boreal-montane (1 species) — Potamogeton subsi- biricus (3) Pacific coastal (2 species) — Rumex beringensis and Salix hookeriana IV. Incompletely circumpolar — 9.7% (7 species) (1) Arctic-alpine (1 species) — Cryptogramma stelleri (2) Boreal-montane (4 species) — Tricophorum pum- ilum and Carex laxa (3) Widespread (2) — Ceratophyllum demersum and Myriophyllum verticillatum V. Circumpolar — 8.3% (6 species) (1) Arctic-alpine (5 species) — Colpodium vahlianum and Cerastium regelii (2) Boreal-montane (1 species) — Viola selkerkii VIL. Amphiatlantic — 1.4% (1 species) — Najas flexilis The main differences between the biogeography of the total flora and the rare flora are the increase in the ratio of Alaska- Yukon endemics in the rare flora which is only 7% of the total flora, and the decrease in the ratio of circumpolar and amphiberingian ele- ments in the rare flora which is 37% and 24% of the total flora respectively. The ecological distribution of the rare flora is: cordilleran (38%), boreal-montane (25%), arctic-alpine (24%), Pacific coastal (6%), temperate disjunct (6%) and widespread (3%). The primary differences in the ecological distribution of the rare flora as compared to the total flora are the increase in the cordilleran element in the rare flora, which is only 12% of the total flora and the decreases in the Pacific coastal and widespread elements in the rare flora which is 12% and 20% of the total flora respectively. Recommendations This inventory has enhanced our understanding of the biodiversity and phytogeography of the region and has filled in the ‘Wrangell Void’ for many species not known to occur in the Park or thought to be rare. 2002 | o OU Alaska-Yukon Percent of Endemic Taxa > Oo ia ~ ad 10 0 NX Tanana White (n=31) (n=22) COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 199 @Amphiberingean WCordilieran WN Pacific Copper (n=37) BT | Chitina (n=37) Gulf (n=24) River Basins FIGURE 3. Ratio of endemic taxa within river basins and Gulf of Alaska, Wrangell-St. Elias National Park and Preserve. However, there are many regions and communities that have yet to be surveyed or need further work. Some of these are: the Gulf of Alaska between Yaku- tat and Icy Bays, the southern St. Elias Mountains and Bagley Icefield, the Tanana lowlands, the White River basin, wetland and aquatic communities throughout the Park, nunataks, areas of high endemisim on the north side of the Park in the Mentasta, Nutzotin and northern Wrangell Mountains, high elevation plateaus in the Jacksina River drainage, and rare plant habitat throughout the Park. Over half of the Park (3.9 million hectares) is designated wilderness, the remaining area (1.4 mil- lion hectares) is theoretically protected. Aircraft, motorboats and cabins are allowed in wilderness in Alaska for subsistence purposes and aircraft conces- sions for climbing, hiking and backpacking are not restricted in the Park. Most of the increase in back- country use is in areas that we have found likely to harbor rare plants. The rate of global changes to ecosystems and plant communities is increasing as are infrastructure development and visitor use in the Park. Therefore, there is a need to survey areas where we anticipate use as well as the unsurveyed regions so that we can begin to assess changes to plant populations and communities. We also need information on the distribution, life history and population ecology of most rare plants so that protection plans may be developed. Notable Collections We documented nine additions to the flora of Alaska, 211 species new to the Park’s flora, 145 range extensions greater than 200 km from previous- ly documented localities and 186 range extensions greater than 60 km from previously documented localities. An annotated list in the phylogenetic order used by William J. Cody in the Flora of the Yukon Territory (1966) is preceded by a synoptic list of the most notable taxa by status. Alaska- Yukon and Park distribution maps for each species follow the annota- tions and are in the order of the annotations. Specimens are housed in the herbarium at Wrangell- St. Elias National Park and Preserve headquarters in Copper Center, Alaska unless otherwise indicated. Synoptic List by Status Taxa new to Alaska (9) Arabis calderi Arabis codyi Arabis drepanoloba Arabis lemmonii Carex petasata Draba lonchocarpa var. thompsonii Festuca minutiflora Najas flexilis Trichophorum pumilum var. rollandii Range Extensions (203) Agoseris glauca Agrostis mertensii Agrostis thurberiana Alopecurus alpinus Aphragmus eschscholtzianus Antennaria media Arabis media Arenaria capillaris Arenaria longipedunculata Arnica amplexicaulis ssp. prima Arnica latifolia Arnica mollis Artemisia hyperborea Aster alpinus ssp. vierhapperi Aster borealis Astragalus adsurgens ssp. viciifolius Astragalus eucosmus ssp. sealei Astragalus harringtonii 200 Astragalus nutzotinensis Astragalus williamsii Botrychium ascendens Botrychium lanceolatum Botrychium minganense Botrychium pinnatum Braya glabella ssp. glabella Braya glabella ssp. purpurascens Callitriche anceps Callitriche hermaphroditica Caltha leptosepala Carex adelostoma Carex albo-nigra Carex buxbaumii Carex chordorrhiza Carex crawfordii Carex eburnea Carex filifolia Carex holostoma Carex interior Carex krausei Carex lasiocarpa ssp. americana Carex laxa Carex lenticularis var. dolia Carex leptalea Carex nardina Carex nigricans Carex obtusata Carex parryana Carex pauciflora Carex petricosa Carex phaeocephala Carex praticola Carex stylosa Carex viridula Carex williamsii Cassiope mertensiana Castilleja chrymactis Castilleja elegans Castilleja yukonis Cerastium regelii Ceratophyllum demersum Chamaerhodos erecta ssp. nutallii Cicuta maculata var. angustifolia Cladothamnus pyrolaeflorus Claytonia tuberosa Collomia linearis Colpodium vahlianum Coptis trifolia Cryptantha shackletteana Cryptogramma crispa Vat. sitchensis Cryptogramma stelleri Cystopteris montana Danthonia intermedia Delphinium brachycentrum Deschampsia brevifolia Douglasia alaskana Douglasia arctica Douglasia gormanii Draba cinerea Draba crassifolia Draba corymbosa Draba densifolia Draba incerta Draba kananaskis THE CANADIAN FIELD-NATURALIST Vol. 116 Draba lactea Draba macounii Draba oligosperma Draba palanderiana Draba porsildii Draba ruaxes Draba stenoloba Draba stenopetala Epilobium lactiflorum Epilobium luteum Erigeron caespitosus Erigeron grandiflorus ssp. arcticus Eriophorum callitrix Eriophorum viridi-carinatum Erysimum pallasii Euphrasia mollis Eutrema edwardsii Fauria crista-galli Festuca brevissima Festuca lenensis Festuca richardsonii Festuca saximontana Gentiana douglasiana Gentiana platypetala Gentianella tenella Glyceria pulchella Gymnocarpium jessoense ssp. parvulum Hackelia deflexa Halimolobos mollis Hippuris montana Impatiens noli-tangere Isoetes echinospora Juncus filiformis Juncus mertensianus Juniperus horizontalis Kobresia sibirica Kobresia simpliciuscula Lesquerella arctica Ligusticum scoticum ssp. hultenii Lupinus kuschei Maianthemum stellatum Minuartia biflora Minuartia dawsonensis Minuartia stricta Mitella pentandra Montia bostockii Myriophyllum verticillatum Nymphaea tetragona ssp. leibergii Osmorhiza depauperata Oxytropis campestris ssp. jordalii Oxytropis huddelsonii Oxytropis scammaniana Papaver alboroseum Papaver radicatum ssp. kluanense Papaver walpolei Penstemon gormanii Phacelia mollis Phippsia algida Phlox hoodii Phlox richardsonii Phyllodoce glanduliflora Plantago eriopoda Poa hispidula Podagrostis aequivalis Polystichum lonchitis 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA Potamogeton friesii Potamogeton pectinatus Potamogeton pusillus var. pusillus Potamogeton praelongus Potamogeton subsibiricus Potamogeton zosteriformis Potentilla arguta ssp. convallaria Potentilla biflora Potentilla diversifolia Potentilla drummondii Potentilla litoralis Potentilla rubricaulis Primula cuneifolia ssp. saxifragifolia Primula egaliksensis Puccinellia deschampsioides Puccinellia interior Ranunculus gelidus ssp. grayi Ranunculus pacificus Ranunculus pedatifidus ssp. affinis Ranunculus sulphureus var. sulphurus Ranunculus trichophyllus var. eradicatus Rumex acestosa ssp. alpestris Rumex beringensis Sagina saginoides Salix commutata Salix rotundifolia ssp. dodgeana Salix setchelliana Salix stolonifera Saussurea angustifolia ssp. yukonensis Saxifraga adscendens ssp. oregonensis Saxifraga bracteata Saxifraga foliolosa Selaginella sibirica Silene involucrata ssp. involucrata Silene menziesii Silene repens Silene uralensis ssp. uralensis Silene williamsii Smelowskia borealis Smelowskia calycina var. porsildii Sparganium minimum Stellaria alaskana Stellaria umbellata Subularia aquatica Swertia perennis Synthyris borealis Taraxacum carneocoloratum Taraxacum phymatocarpum Thlaspi arcticum Trisetum sibiricum ssp. litorale Vahlodea atropurpurea Veronica serpyllifolia ssp. humifusa Viola adunca Viola biflora Viola selkirkii Alaska Natural Heritage Program State Rare Plants (State Rank < = 3) (69) Agoseris glauca Agrostis thurberiana Aphragmus eschscholtzianus Arabis calderi Arabis codyi Arabis drepanoloba Arabis lemmonii Arenaria longipedunculata Arnica mollis Astragalus harringtonii Botrychium ascendens Carex adelostoma Carex crawfordii Carex eburnea Carex holostoma Carex interior Carex laxa Carex lenticularis var. dolia Carex parryana Carex petasata Cerastium regelii Ceratophyllum demersum Colpodium vahlianum Cryptantha shackletteana Cryptogramma stelleri Cystopteris montana Douglasia alaskana Douglasia arctica Douglasia gormanii Draba densifolia Draba incerta Draba kananaskis Draba lonchocarpa var. thompsonii Draba porsildii Draba ruaxes Draba stenopetala Eriophorum viridi-carinatum Erysimum pallasii var. pallasii Festuca lenensis Festuca minutiflora Glyceria pulchella Juniperus horizontalis Lupinus kuschei Maianthemum stellatum Minuartia biflora Montia bostockii Myriophyllum verticillatum Najas flexilis Oxytropis huddelsonii Papaver alboroseum Papaver walpolei Phacelia mollis Phlox hoodii Phlox sibirica ssp. richardsonii Potamogeton subsibiricus Potentilla drummondii Potentilla rubricaulis Ranunculus pacificus Rumex beringensis Salix setchelliana Saxifraga adscendens ssp. oregonensis Smelowskia calycina var. porsildii Stellaria alaskana Stellaria umbellata Taraxacum carneocoloratum Thlaspi arcticum Trichophorum pumilum var. rollandii Trisetum sibiricum ssp. litorale Viola selkirkii 201 202 Species Accounts SELAGINELLACEAE Selaginella sibirica (Milde) Hieron, Siberian Spike- Moss — CHUGACH MOUNTAINS: rare in tundra, Granitic Creek, 1768 m, 61°6.19’ N 142°55.03’ W, M. Cook 94220A, 8 July 1994. NUTZOTIN MOUNTAINS: scattered in rock crevices, Lick Ridge, 1554 m, 62°28.44’ N 142°15.06' W, M. Cook 95118, 29 June 1995. WRANGELL MOUNTAINS: scat- tered on SE-facing bluff, Nabesna River, 1106 m, 62°15.05’ N 142°54.68’ W, M. Cook 96189, C. Roland 96-161, 19 June 1996; outcrop crevices, Nikolai Mine, 1695 m, 61°27’ N 142°39' W, Batten & Barker 96-055, 24 July 1996; outcrop, Nikolai Ridge, 1372 m, 61°26.5'’ N 142°43' W, Batten & Barker 96-157, 26 July 1996 (ALA); rock ledges, Crystalline Hills, 1585 m, 61°23.59'°N 143°31.85' W, M. Cook 95276, 19 July 1995; in moss under open willow scrub, Fish Creek, 1067 m, 62°16.92' N 142°58.99' W, M. Cook 95312, 27 July 1995. This amphiberingean species occurs in dry, rocky situations throughout northern Alaska. The Granitic Creek locality extends its range 272 km to the southeast into the Chugach Mountains from a station in the Alaska Range reported by Hultén (1968). It was also collected at several intermediate localities in the Wrangell and Nutzotin Mountains. Map 1. ISOETACEAE Isoetes echinospora Durieu (J. muricata Durieu), Quillwort — WRANGELL MOUNTAINS: abundant, uprooted and lining shoreline, Lake 2870, 0.2 km N of Tanada Creek, 875 m, 62°31.38' N 143° 28.12’ W, C. Roland 95-096, 28 June 1995; few clumped in muck, 0.6 m deep water, Fox Farm Lake, 727 m, 62°19.98’ N 144°50.02’ W, M. Cook 95308, 25 July 1995; abundant submerged in 30 cm deep water, Lake 2990’, 3.6 km west of Copper Lake, 911 m, 62°24.53' N 143°42.68' W, C. Roland 96-913, 11 August 1996. This boreal North American aquatic species has been found at a few widely separated sites across central Alaska and the Yukon Territory. The specimens cited above extend its range 80 km southeast into the Wrangell Mountains from a station in the Alaska Range (Hultén 1968). Map 2. OPHIOGLOSSACEAE Botrychium ascendens W. H. Wagner, Triangle- Lobe Moonwort — NUTZOTIN MOUNTAINS: few on S-facing scree slope, Gold Hill, 1434 m, M. Cook 3520B, 18 July 1999. This North American species with a cordilleran distribu- tion was known from two localities in Alaska and one in the Yukon Territory (Cody 1994). It is rare in Alaska (G3? S1) and Cody (1994) suggested that it be added to the list of rare species for the Yukon Territory. The locality cited above extends its range 266 km to the southwest of a station in the Yukon Territory and 356 km northwest of a collection locality in the Yakutat Quad (Akwe Dunes, 59°21.78' N 138°50.45’ W, M.C. Stensvold 7304, 27 June 1996 (ALA)). Map 3. THE CANADIAN FIELD-NATURALIST Vol. 116 Botrychium lanceolatum (Gmel.) Angstr., Lanceleaf Grapefern — CHUGACH MOUNTAINS: rare on S-fac- ing slopes, Juniper Island, 1346 m, 60°36.17’ N 142°14.96’ W, C. Roland 94-208, 10 July 1994; openings in willow thicket, Granite Creek, 823 m, 60°44’ N 142°13’ W, Batten & Barker 96-354B, 30 July 1996 (ALA); open low shrub birch scrub, Granite Creek, 579 m, 60°44’ N 142°32’ W, Parker & Duffy 6718, 6 August 1996. WRANGELL MOouN- TAINS: earth-flow scar on steep S-facing slope, Nikolai Pass, 1372 m, 61°26.5’ N 142°43’ W, Batten & Barker 96-174B, 26 July 1996 (ALA); scattered in grassy gully, Long Glacier, 1399 m, 61°48.04' N 144°6.27' W, M. Cook 96702, C. Roland 96-851, 6 August 1996. This moonwort occurs in maritime-influenced boreal regions and scattered inland locations throughout the cir- cumpolar north. In Alaska, it occurs most frequently along the southern coast from the panhandle through the Aleutian Islands. Our collections extend its range 175 km northeast into the Wrangell and Chugach Mountains from a collec- tion in the Cordova Quad (60°58’ N 145° W, C. L. Parker 1830B, 12 August 1986 (ALA)) and connect the distribution 202 km to the east in the Yukon Territory (Cody 1996). Map 4. Botrychium minganense Victorin, Mingan Moon- wort — NUTZOTIN MOUNTAINS: few in Dryas- graminoid-forb herbaceous stringer at base of scree slope, headwaters of Alder Creek, 1554 m, 62°28.44’ N 142°15.06’ W, M. Cook 95139, 30 June 1995; open shrub birch-willow scrub, Ptarmigan Lake, 1128 m, 61°50.12' N 141°9.15' W, Duffy & Barnes 96-296, 8 August 1996; few on S-facing scree slope, Gold Hill, 1434 m, M. Cook 3520C, 18 July 1999. This moonwort occurs in scattered localities across bore- al North America and as far south as Arizona in the western cordillera. It is known from only two other localities in Alaska, 405 km to the northwest in the Healy Quad (Chulitna River, 62°59.75’ N 150°6.83’ W, C. Roland 3493B, 18 August 1998 (ALA)) and 308 km to the south- east in the Yakutat Quad (59°19.35’ N 138°40.14’ W, M. Stensvold 7313, 16 July 1998 (ALA)). Map 5. Botrychium pinnatum H. St. John (B. boreale (E. Fries) Milde), Northeastern Moonwort — CHUGACH MOUNTAINS: scrub thicket, Granite Creek, 1067 m, 61°5.46’ N 142°54.05’ W, J. Bolivar 84-172, 7 August 1984; rare in disturbed mineral soil, upper Golchonda Creek, 1402 m, 61°2.89' N 143°24.42' W, M. Cook 3200, 28 July 1998; openings in willow thicket, Granite River, 823 m, 60°44’ N 142°13’ W, Batten & Barker 96-354C, 30 July 1996. NUTZOTIN MOunrAINS: few in Dryas-graminoid tundra, head- waters of Alder Creek, 1554 m, 62°28.44’ N 142°15.06'’ W, M. Cook 95121, 29 June 1995. WRANGELL MOUNTAINS: scattered in meadow, vic. Chitistone Falls, 1177 m, 61°32.27’ N 142°11.49’ W, C. Roland 96-632, 22 July 1996; steep E-facing meadow, Nikolai Mine, 1695 m, 61°27’ N 142°39’ W, Batten & Barker 96-074B, 24 July 1996 (ALA); 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 203 I. Selaginella sibirica 4. Botrychium lanceolatum 204 THE CANADIAN FIELD-NATURALIST Vol. 116 8. Cryptogramma stelleri 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 205 11. Polystichum lonchitis 12. Juniperus horizontalis 206 earthflow scar on steep slope, Nikolai Pass, 1372 m, 61°26.5’ N 142°43’ W, Batten & Barker 96-174A, 26 July 1996; moist silt near stream, W slope of Chitistone Mountain, 1219 m, 61°28’ N 142°33' W, Batten & Barker 96-146, 26 July 1996 (ALA); patchy in bare area, vic. Long Glacier, 1399 m, 61°48.04' N 144°6.27' W, M. Cook 96701, 96703, 96716, 8 August 1996; few in mesic forb herbaceous vegetation, vic. Grant Creek, 1250 m, 61°17.65' N 143°56.34' W, M. Cook 96367, 7 July 1996. This amphiberingean species occurs in scattered locali- ties throughout boreal areas of North America. It was considered rare in the Yukon Territory by Douglas et al. in 1981. The collections cited above extend its range 324 km east into the Wrangell Mountains, 341 km to the northeast into the Nutzotin Mountains and 367 km east into the Chugach Mountains from a collection in the Anchorage Quad (61°50.0' N 149°18.0’ W, Parker & Murray 2097, 10 August 1989 (ALA)). These new localities also connect the range to the west with the range 256 km to the east in the Yukon Territory. Map 6. PTERIDACEAE Cryptogramma crispa (L.) R. Br. var. sitchensis (Rupr.) C. Christens., Alaska Parsley Fern — CHUGACH MOUNTAINS: rubble slope, Granite Creek, 579 m, 60°44’ N 142°32’ W, Parker & Duffy 6709, 6 August 1996; scattered in moss, Lake Creek, 975 m, 61°11.4’ N 143°49.45’ W, M. Cook 96401, 13 July 1996; scree, Hanagita Peak, 1186 m, 61°4.91' N 143°38.86’ W, M. Cook 96421, 14 July 1996; occasional on lateral moraine, 12 Mile Basin, 1271 m, 60°48.85’ N 142°33.52’ W, M. Cook 96636, 30 July 1996; dry boulder area, Upper Tebay Lake, 610 m, 61°11.5’ N 144°23.5’ W, L. A. & E. G. Viereck 11083, 7 July 1996. St. ELIAS MOUNTAINS: closed tall alder scrub on glacial rubble, Atrevida Glacier, 59°57.06' N 139°48.5’ W, B. Haller s.n., 19 August 1987. This fern variety is endemic to Alaska, the Yukon Territory, northern British Columbia and the District of Mackenzie, Northwest Territories where it grows in moist talus and rock crevices. The specimens cited above extend its range 293 km northeast into the Chugach Mountains from a station near Seward (Hultén 1968) and connect the range 200 km to the southeast at Yakutat (Hultén 1968). Map 7. Cryptogramma stelleri (S.G. Gmel.) Prantl., Fragile Rock-Brake — NUTZOTIN MOUNTAINS: head of Sheep Creek, 6 km above confluence with Chisana River, 1067 m, 62°7.95' N 141°54.9’ W, J. Bolivar 84-82, 26 June 1984; rare under boulders on SW- facing slope, Carl Creek, 1920 m, 62°3.52' N 141°36.27' W, M. Cook 95006, 14 June 1994; rare in rock crevices, Carden Hills, 1311 m, 62°18.44’ N 141°11.55’ W, M. Cook 94166, 25 June 1994. WRANGELL MOUNTAINS: rare in rock crevices, Monte Cristo Creek, 975 m, 62°13.66’ N 142°56.35’ W, C. Roland 96-147, 96-149, 18 June 1996; rare under boulders, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24’ W, C. Roland 95-133A, 4 July 1995. THE CANADIAN FIELD-NATURALIST Vol. 116 This fern is circumpolar and widespread but rare throughout its distribution. It is considered rare in both Alaska (G5 $2S3) and the Yukon Territory (Douglas et al. 1981). Our collections extend its range 312 km south into the Nutzotin and Wrangell Mountains from a collection in the Big Delta Quad (Brigadier Road, 64°40.75' N 146°12.38' W, Duffy & Lipkin 95-654, 12 July 1995 (ALA)). Map 8. DRYOPTERIDACEAE : Cystopteris montana (Lam.) Bernh., Mountain Fragile Fern — CHUGACH MOounrtaAINs: SE-facing bench, Lower Bremner River, 457 m, 61°3.39’ N 144°26.8' W, K.A. Teare 1809, 20 July 1984. MENTASTA MOUNTAINS: scattered at forest margin toe of talus slope, Totschunda Creek, 1280 m, 62°27.85' N 142°40.68'’ W, C. Roland 96-298, 24 June 1996. WRANGELL MOUNTAINS: limestone rocks in alder thicket, Fish Creek, 1067 m, 62°16.92’ N 142°58.99' W, C. Roland 95-293, 29 July 1995; scattered in moss-forb tundra on limestone outcrop, Lakina Glacier, 1219 m, 61°33.64’ N 143°19.01’ W, C. Roland 96-689, M. Cook 96534, 24 July 1996. This fern is circumboreal and is considered rare in Alaska (G4S3) where it occurs in montane meadows and thickets, often on calcareous substrates. It had been collect- ed in the Wrangell Mountains (Bonanza Ridge, 61°30’ N 142°51' W, Nordell & Schmitt 580, 1976 (ALA)). The new collections cited above extend its range 114 km north into the Mentasta Mountains and 100 km south into the Chugach Mountains. Map 9. Gymnocarpium jessoense (Koidz.) Koidz. ssp. parvu- lum Sarvela (G. robertianum (Hoffm.) Newm.), Limestone Oak-Fern — NUTZOTIN MOUNTAINS: rare on rock ledges under low birch/willow scrub, Carden Hills, 1311 m, 62°18.44’ N 141°11.55' W, M. Cook 94173, 25 June 1994. This species occurs in boreal areas throughout the cir- cumpolar region except Greenland. In Alaska and the Yukon it is generally restricted to the central Yukon River drainage except for a single locality near Anchorage report- ed by Hultén (1968). It is considered rare in the Yukon Territory (Douglas et al. 1981). The specimen cited above extends its range 180 km south into the Nutzotin Mountains from a station near Chicken in the Yukon-Tanana uplands (Hultén 1968). Map 10. Polystichum lonchitis (L.) Roth, Holly Fern — CHUGACH MOUNTAINS: rare on outcrops, E Fork Little Bremner River, 1219 m, 61°4.53’ N 144°3.6' W, L. A. & E. G. Viereck 11104, 7 July 1996; rare on E-facing rocks of ridge, vic. Spirit Mountain, 1204 m, 61°21.09’ N 144°32.4’ W, C. Roland 96- 624, 17 July 1996; scattered on S-facing talus slope, vic. 12-mile Creek, Granite Range, 1335 m, 60°49.71' N 142°33.34' W, C. Roland 96-813, 30 July 1996. GULF OF ALASKA: moraine, Chaix Hills, Icy Bay, 427 m, 60°7.52’ N 141°8.1’ W, B. Haller s.n., 24 August 1987. Map 11. This fern is circumpolar with a boreal-montane distribu- tion and is considered rare in the Yukon Territory (Douglas 2002 et al. 1981). Our collections extend its range 275 km south- east into the southern St. Elias Mountains from a station near Thompson Pass (61°08’ N 145° 44’ W, C. L. Parker 2462, 23 July 1990 (ALA)) and connect the distribution 110 km to the southeast at Yakutat (F. Funston 126, 15 August 1892 (US), Hultén 1941). CUPRESSACEAE Juniperus horizontalis Moench, Creeping Savin — CHUGACH MOUNTAINS: S-facing bank of lake, vic. East Fork Kiagna River, Granite Range, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-202, 27 July 1996. St. ELIAS MOUNTAINS: sandy hillside, Hubert’s Landing, Chitina River, 671 m, 61°2.7' N 141°38’ W, D. Miquelle 84-38, 15 July 1984; allu- vial floodplain of Clear Stream, Chitina River, 488 m, 61°5.82'’ N 141°57.92’ W, Duffy & Barnes 96-005, 8 August 1996. WRANGELL MOUNTAINS: small patch in gully within steppe, Dadina River bluff, Copper River, 393 m, 61°51.49’ N 145°0.72' W, C. Roland 96-304, 1 July 1996; scattered on S- facing bluff in aspen woodland, Crystalline Hills, 602 m, 61°23.12' N 143°36.16’ W, M. Cook 3129, 15 June 1998 (ALA). This species occurs in dry situations across boreal North America but is considered rare in Alaska (G5 $1S2) where it is known only from the Copper River drainage, the vicin- ity of Anchorage and a single locality in the Tanana River valley. It is common on bluffs along the Chitina and middle Copper River drainages where it is often found growing with Juniperus communis L. It had been collected in the Wrangell Mountains (Bonanza Ridge, 61°30’ N 142°51’ W, Nordell & Schmitt s.n., 1976 (LD & ALA), Nordell and Schmitt (1977) and at the head of the Chitina River (H. M. Laing 10, 11 (CAN), Hultén (1941) and Porsild (1939). Our collections extend its range 25 km north into the Copper River valley. Map 12. SPARGANIACEAE Sparganium minimum (Hartm.) E. Fries (S. natans L.), Arctic Burr-Reed — CHUGACH MOUNTAINS: pond margin, Middle Fork of the Bremner River, 869 m, 60°55.05’ N 143°43.86' W, Duffy & Barnes 96-114, 6 August 1996; mud of small dried pond, Tana River Flats, 442 m, 61°14.45’ N 142°57.21' W, Duffy & Barnes 96-135, 7 August 1996; scattered in pond, upper Tebay Lake, 579 m, 61°11’ N 144°24’ W, Parker & Gracz 6767, 7 August 1996. WRANGELL MOUNTAINS: shallow water, vic. Tanada Creek, 927 m, 62°30.34’ N 143°25.44’ W, Moran & Roland 95-18, 29 June 1995. This species is circumboreal and is considered rare in the Yukon Territory (Douglas et al. 1981). The specimens cited above extend its range 130 km south into the Wrangell Mountains from a station near Paxson (Hultén 1968) and 280 km east into the Chugach Mountains from a collection in the Anchorage Quad (Otter Lake, 61°17.53’ N 149°44.17' W, Duffy & Tande 1020, 3 August 1994 (ALA)). Map 13. POTAMOGETONACEAE Potamogeton friesii Rupr., Fries’ Pondweed — WRANGELL MOUNTAINS: abundant in small, rock- COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 207 floored pond in 30-100 cm of water, Fish Creek, 1067 m, 62°16.92’ N 142°58.99' W, C. Roland 95- 286A, 29 July 1995. This circumpolar species is widespread but with wide gaps. The specimen cited above extends its range 163 km northwest into the Wrangell Mountains from a station near Tonsina (Hultén 1968) Map 14. Potamogeton pectinatus L., Sago Pondweed — COPPER RIVER BASIN: floating next to shore in pond, Old Edgerton Highway, km 2.5, 415 m, 61°49.3’ N 145°10.7' W, M. Cook 95372, 5 August 1995. This locality extends the range of this pondweed 106 km east into the Copper River basin from a station in the Anchorage Quad (McRoberts Creek, 61°33.0’ N 148°55.0’ W, Batten & Reed 80-119A, 1 July 1980 (ALA)). Map 15 Potamogeton praelongus Wulf., White-Stemmed Pondweed — WRANGELL MOUNTAINS: Scattered in shallow water around pond, vic. Orange Hill, 884 m, 62°12.63'’ N 142°52.28’ W, C. Roland 96-121B, 17 June 1996; shore of shallow lake, vic. Monte Cristo Creek, 1021 m, 62°14.07' N 142°56.85’ W, C. Roland 96-141, 18 June 1996; common in deep water of pond, vic. of Copper Lake, 911 m, 62°24.53’ N 143°42.68' W, C. Roland 96-908, 11 August 1996; common in deep water of lake, vic. Lake 2910, Copper River, 823 m, 62°26.35’ N 143°40.77' W, C. Roland 96-926, 12 August 1996. CHUGACH MOUNTAINS: fresh pondweed shallows, East Flowers Lake, 404 m, 61°5.5' N 142°34.2' W, Duffy & Barnes 96-205, 8 August 1996; 1 m deep water, Chokosna Lake, 619 m, 61°27.45’ N 143°49’ W, C. Roland 94-328A, 10 August 1994; rooted in organic muck, Lakina Pond, 792 m, 61°29.49’ N 143°25.6' W, C. Roland 96-677, 24 July 1996. This pondweed, incompletely circumpolar and widespread in its distribution, is rare in the Yukon Territory (Douglas et al. 1981). Our collections extend its range 199 km south into the Chugach Mountains from a collection near Chistochina (62°30.0' N 144°50.0' W, G. Smith 2302, 16 May 1954 (ALA)). Map 16. Potamogeton pusillus L. var. pusillus (P. berchtoldti Fieber), Small Pondweed — CHUGACH MOUNTAINS: mixed freshwater herbaceous, pond margin | m deep, Tana Flats, 442 m, 61°14.45’ N, 142°57.21' W, Duffy & Barnes 96-130B, 96-127, 96-122B, 7 August 1996; floating adjacent to shore, Old Edgerton pond, 415 m, 61°49.3’ N 145°10.7’ W, M. Cook 95374, 5 August 1995; common in shallow water, Happel Slough, 180 m, 61°2.13’ N 144°29.43' W, C. Roland 96-618, 17 July 1996. WRANGELL MOUNTAINS: beaver pond E of Chititu Creek, 457 m, 61°22.16' N 142°40.29' W, M. Duffy 91043, 8 July 1991; in 1 m deep water, Chokosna Lake, 619 m, 61°27.45' N 143°49’ W, C. Roland 94-328C, 10 August 1994; common, submerged in shallow water, vic. Indian Creek, 655 m, 62°38.1' N 144°20.2' W, M. Cook 95343, 1 August 1995; rare on bottom in 30 cm deep water, Lake 2990’, Copper River, 911 m, 62°24.53' 208 THE CANADIAN FIELD-NATURALIST Vol. 116 16. Potamogeton praelongus 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 209 20. Najas flexilis 210 N 143°42.68’ W, C. Roland 96-909, 11 August 1996; common rooted in muddy bottom, Copper River pond 0.5 mi E of Lake 2910’, 823 m, 62°26.35’ N 143°40.77' W, C. Roland 96-925, 12 August 1996; forming dense stands throughout bottom of lake, Fish Creek, 1067 m, 62°16.92’ N 142°58.99’ W, C. Roland 95-273, 28 July 1995, C. Roland 95- 272, M. Cook 95321, 29 July 1995. This species is circumpolar with a widespread distribu- tion. The specimens cited above extend its range 168 km southeast into the Wrangell Mountains from a station near Paxson (Hultén 1968) and 99 km east into the Chugach Mountains from a station near Thompson Pass (Hultén 1968). Map 17. Potamogeton subsibiricus Hagstr., Yenisei River Pondweed — WRANGELL MOUNTAINS: rare in shal- low water, Lake 2990’, upper Copper River Basin, 911 m, 62°24.53' N 143°42.68’ W, C. Roland 96- 919, 11 August 1996. This amphiberingean species occurs in widely separated localities in boreal regions of Alaska and the Yukon. It is considered rare in Alaska (G3 S3) and the Yukon Territory (Douglas et al. 1981). The Copper River locality extends its range 87 km east into the Wrangell Mountains from a station along the Richardson Highway between Gakona and Paxson (Hultén 1968). Map 18. Potamogeton zosteriformis Fern., Flatstem Pondweed — CHUGACH MOUNTAINS: common just below surface next to shore, pond at mile 1.3 Old Edgerton Highway, 415 m, 61°49.3’ N 145°10.7' W, M. Cook 95375, 5 August 1995. This North American boreal-montane species is of limt- ed distribution in the Yukon Territory (Cody 1994; Cody et al. 1998). The specimen cited above extends its range 188 km to the northwest into the Copper River basin from a locality in the Anchorage Quad (Little Kiowa Lake, 61°15.5’ N 149°39.38' W, Duffy & Tande 1031, 4 August 1994 (ALA)). Map 19. NAJADACEAE Najas flexilis (Willdenow) Rostkov. & Schmidt, Wavy Waternymph — WRANGELL MOUNTAINS: shal- low water, Chokosna Lake, 619 m, 61°27.45’ N 143°49' W, C. Roland 94-325, 10 August 1994. This species is new to the flora of Alaska and although it was collected in 1986 in the Fort Yukon Quad (Heglund Plot Lake 23, & Lake 522 near Preacher Creek, 66°01'N 144° 42’ W, P. Heglund 86-351, 86-363, 16 & 17 July 1986 (ALA)), it was not reported. It is now known from a third locality in Alaska in the Anchorage Quad (Little Kiowa Lake, 61°15.48’ N 149°39.38' W, Duffy & Tande 1030, 4 August 1994). The Wrangell Mountains locality is 243 km northeast of the Anchorage Quad collection site. This species has an amphiatlantic distribution, and is widespread across North America and Europe. It is now considered rare in Alaska (G5 $182). Map 20. POACEAE Agrostis mertensii Trin. (A. borealis Hartm.), Red Bentgrass — CHUGACH MOUNTAINS: scattered in meadow on S-facing slope, vic. Twelve-Mile Creek, 1335 m, 60°49.71’ N 142°33.34’ W, C. Roland 96- THE CANADIAN FIELD-NATURALIST Vol. 116 808, 30 July 1996; scattered in subarctic lowland sedge wet meadow, upper Tebay Lake, 579 m, 61°11’ N 144°24’ W, Parker & Gracao7se. 7 August 1996. WRANGELL MOUNTAINS: scattered in lichen-heath tundra, plateau vic. Long Glacier, 1399 m, 61°48.04' N 144°6.27’ W, C. Roland 96-853, 96-870, 6 August 1996. This species is amphiberingean with an arctic-alpine distribution. The specimens cited above extend the range 95 km south into the Chugach Mountains from a collection on Bonanza Peak in the southern Wrangell Mountains (61°30' N 142°51' W, Nordell & Schmitt 490, 1976 (LD & ALA), Nordell & Schmitt s.n., 1978). The new localities connect the range 106 km to the west in the Anchorage Quad (Brilliant Glacier, 61°06’ N 147°29’ W, M. Duffy s.n., 19 August 1993 (ALA)) with a locality 441 km to the southeast near Skagway (59°33.0’ N 135°06.0’ W, A.R. Batten 88-516, 30 August 1988 (ALA)). Map 21. Agrostis thurberiana A. S. Hitche. (Podagrostis thurberiana (Hitchc.) Hult.), Thurber’s Bentgrass — CHUGACH MOUNTAINS: herbaceous meadow, under Salix barclayi, vic. 12-mile Creek, 1335 m, 60°49.71’ N 142°33.34’ W, C. Roland 96-833, 7 July 1996. GULF OF ALASKA: tall forb herbaceous slope, Amphitheather Knob, 427 m, 59°57’ N 139°46' W, M. Cook 87-165, 19 August 1987. This western North American species of Bentgrass occurs in scatterd localities along the southern coast of Alaska where it is considered rare (G5 $2). Our collections connect its range 212 km to the southwest in the Seward Quad (Iron Mountain, 60°22.00' N 147°39.0' W, M. Duffy 93-1095, 17 August 1993 (ALA)) with a station near Pelican 345 km to the southeast (Hultén 1968). Map 22. Alopecurus alpinus J.E. Smith (A. borealis Trin.), Mountain Foxtail — NUTZOTIN MOUNTAINS: abun- dant in mesic tundra on SW-facing slopes, Wiki Creek, 1411 m, 61°54.46’ N 141°10.68" W, C. Roland 96-088, 10 June 1996; disturbed wet sedge meadow, Francis Creek, 1085 m, 61°51.93’ N 141°9.4’ W, Duffy & Barnes 96-233, 8 August 1996. WRANGELL MOUNTAINS: common in bare clay soil of frost boils, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24”" W, C. Roland 95-101, 30 June 1995; Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5’ W, M. Potkin 95-115, 95-142, 31 July 1995. The collections cited above extend the range of this circumpolar, arctic-alpine grass south 182 km into the Wrangell Mountains and 244 km into the Nutzotin Mountains from a station in the Tanacross Quad (W Fork of the Dennison, 63°50.00' N 142°15.0’ W, Smith & Viereck 2395, 24 June 1954 (ALA)). Map 23. Colpodium vahlianum (Liebm.) Nevski (Puccinellia vahliana (Liebm.) Scribn. & Merr.), Vahl’s Alkalai Grass — WRANGELL MOUNTAINS: scattered in wet frost boils on limestone ridge, Cooper Pass, 1942 m, 62°17.16' N 142°31.44’ W, C. Roland 94-279, 24 July 1994 (ALA). This arctic—alpine tundra grass is circumpolar and is considered rare in both Alaska (G4 S3) and the Yukon 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 24. Colpodium vahlianum 211 | wr 8 ee eee ed Territory (Douglas et al. 1981). The collection cited above is 719 km south of the nearest locality on the arctic coast (Table Mountain Quad: Double Mt., 68°44.00’ N 143°35.0' W, A.R. Batten 75-514a, 22 July 1975 (ALA)) and 318 km west of a station in the Yukon Territory near Haines Junction (Cody 1996). Map 24. Danthonia intermedia Vasey, Timber Wild Oat Grass — CHUGACH MOUNTAINS: abundant in turfy meadow on SE-facing slope, Granite Creek, 1829 m, 61°0.22' N 141° 51.1’ W, C. Roland 96-752, 28 July 1996; abundant among heath hummocks on S-facing slope, vic 12-mile Creek, 1335 m, 60°49.71’ N 142°33.34’ W, C. Roland 96-811, 30 July 1996; occasional in open patches of birch scrub, Granite Creek, 884 m, 60°44.63' N 142°6.05’ W, M. Cook 96604, C. Roland 96-792, 29 July 1996; moist herbaceous bank, Short River Pond, 503 m, 61°5.35’ N 141°56.3’ W, Parker & Duffy 6687, 6 August 1996; scattered in subarctic lowland sedge wet meadow, Falls Creek, 579 m, 61°11’ N 144°24’ W, Parker & Gracz 6760, 7 August 1996; mixed mesic forb herbaceous mead- ow & dry S-facing slope, Middle Fork of the Bremner River, 869 m, 60°55.05’ N 143°43.86' W, Duffy & Barnes 96-087, 96-104, 8 August 1996. ST. ELIAS MOUNTAINS: locally common in graminoid meadow, Blondie Ridge, 1951 m, 60°53.94’ N 141° 6.75’ W, C. Roland 95-171, 12 July 1995. WRANGELL MOUNTAINS: dominant in dry graminoid herbaceous meadow, Cheshnina Plateau, 1399 m, 61°48.04’ N 144°6.27' W, M. Cook 96715, C. Roland 96-848, 96- 864, 5 August 1996; scattered in turfy meadow, Chitistone Falls, 1177 m, 61°32.27' N 142°11.49’ W, C. Roland 96-635, 22 July 1996. This species occurs in boreal meadows across North America and in Kamchatka (Hultén 1968). In Alaska, it was reported from the Kenai Peninsula and near Anchorage by Hultén (1968). Our specimens extend its range northeast 163 km into the Wrangell Mountains, 239 km into the St. Elias Mountains and 182 km into the Chugach Mountains from a collection near Cordova (60°30.93’ N 145°25.04’ W, C. Parker 1944, 15 August 1986 (ALA)) and connect the range 215 km to the east in the Yukon Territory near Haines Junction (Cody 1996). Map 25. Deschampsia brevifolia R. Br. (D. caespitosa ssp. brevifolia (R. Br.) Tzvelev), Tufted Hairgrass — WRANGELL MOUNTAINS: patchy in wet graminoid tundra, vic. Sheep Glacier, 1478 m, 62°21.85' N 144°23.73' W, M. Cook 94425, 30 July 1994 (ALA); Lakes Plateau, 1890 m, 62°4.4' N 143°23.5’ W, M. Potkin 95-089, 29 July 1995; few on gravel bar, Copper River near Lake 2990’, 911 m, 62°24.53’ N 143°42.68' W, M. Cook 96737, 11 August 1996; scattered in openings between alders on gravel bar, vic. Batzulnetas, 640 m, 62°38.6’ N 143°52.09’ W, M. Cook 96749, 96752, 13 August 1996. The specimens cited above extend the range of this circumpolar, arctic-alpine species 291 km south into the Wrangell Mountains from a station in the Alaska Range (Hultén 1968) and connect the range 180 km to the east in the Yukon Territory (Cody 1996). Map 26. THE CANADIAN FIELD-NATURALIST Vol. 116 Festuca brevissima Jurtzev, Alaska Fescue — CHUGACH MOUNTAINS: scattered along ridge between Chakina River and Granitic Creek, 1768 m, 61°6.19’ N 142°55.03’ W, M. Cook 94227A, 7 July 1994. MENTASTA MOUNTAINS: bare soil of exposed creek cutbank, Lost Creek, 1722°m;.,.6274638 5 sy 143°12.03' W, Beck & Cook 95240, 4 July 1995. NUTZOTIN MOUNTAINS: scattered in orange rhyolite scree, plateau between Bryan and Willow Creeks, 1829 m, 61°58.97' N 141°50.03’ W, M. Cook 95052, 18 June 1995; occasional in moss stringers on unsta- ble granitic rubble slope, Antler Creek, 1585 m, 62°25.78’ N 142°22.11' W, Cook & Beck 95103, 27 June 1995; scattered in bare mineral soil on top of rock outcrop, alpine basin at head of Lick Creek, 1554 m, 62°28.44’ N 142°15.06’ W, M. Cook 95125, 29 June 1995; scattered on sheep trail along ridge, Carl Creek, 1554 m, 62°2.81' N 141°34.65’ W, M. Cook 3146A, 1 July 1998 (ALA); scattered in moist gravel, Wiki Basin, 1585 m, 61°55.08’ N 141°12.07’ W, M. Cook 3178, 15 July 1998 (ALA). WRANGELL MOUNTAINS: common in dry gravel, NE slope of Mt. Drum, 1433 m, 62°8.83' N 144°30.18’ W, C. Roland 94-070, 11 June 1994; occasional on S-facing basalt outcrops, Cone Ridge; 2073 m, 6276227 a 143°18.47' W, M. Cook 94388, 26 July 1994; occa- sional in unstable S-facing rubble and cinders, Cone Ridge, 2073 m, 62°8.21' N 143°18.47' W, C. Roland 94-317, 27 July 1994; scattered in Cassiope- graminoid tundra on SW-facing slope, Chetaslina plateau, 1615 m, 61°56.51’ N 144°25.93’ W, M. Cook 94478, 16 August 1994; scattered in gravel on frost-boils, Jaegar Mesa, 1893 m, 62°15.9' N 143°1.24’ W, C. Roland 95-110, 95-114, 95-119, 95- 131, 1 & 2 July 1995; scattered in gravel patches amongst Dryas tundra, Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85' W, M. Cook 95266, 18 July 1995; Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5’ W, M. Potkin 95-090, 29 July 1995; moist bare min- eral soil of solifluction lobes, Black Mountain, 1481 m, 62°20.85’ N 143°44.9' W, M. Cook 95141, 95159, 7 July 1995; scattered in bare gravel patches within Cassiope-ericaceous heath, Boulder Lake, 1036 m, 62°31.27' N 144°11.26’ W, M. Cook 95357, 8 August 1995, confirmed by Signe Fredericksen (C) 1998; occasional in morainal deposits, valley between Ruddy Mountain and Mt. Drum, 1615 m, 62°4.61' N 144°46.39’ W, C. Roland 96-326, 2 July 1996; N-facing scree, Nikolai Pass, 1280 m, 61°26’ N 142°40' W, Batten & Barker 96-008, 23 July 1996; bouldery dry tundra on south slope, Nikolai Pass, 1280 m, 61°26’ N 142°40’ W, Batten & Barker 96-100, 24 July 1996 (ALA); scattered in bare, . gravel-sized rhyolite scree on SE-facing slope, Kuskulana Pass, 1545 m, 61°33.72’ N 143°39.7' W, C. Roland 96-709, 26 July 1996; occasional in bar- ren, gravelly sites on SW-facing slope, Snyder Peak, 1524 m, 62°4.47' N 144°30.51' W, C. Roland 96- 380, 5 July 1996. 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 213 28. Festuca lenensis 214 This amphiberingean arctic-alpine species is rare in Alaska (G4 S$3S4). The specimens cited above extend its range 265 km into the Chugach Mountains from a collec- tion in the Alaska Range (Independent Ridge, 63°40.0’ N 144°54.0’ W, L.A. Spetzman s.n., 20 June 1957 (ALA)). It is fairly common at high elevations in the northern region of the Park, where it was collected at numerous localities in the Mentasta, Nutzotin and Wrangell mountains. Map 27. Festuca lenensis Drobov (F. ovina L. ssp. alaskensis Holmen), Tundra Fescue — MENTASTA MOUNTAINS: common in gravelly sites on limestone ridge in dry Dryas octopetala tundra, Lost Creek, 1646 m, 62°34.58'’ N 143°5.58’ W, C. Roland 94-268B, 23 July 1994. NUTZOTIN MOUNTAINS: scattered on unstable S-facing scree slope, Horsfeld Creek, 1768 m, 62°2.88' N 141°13.18’ W, M. Cook 94152, C. Roland 94-129C, 24 June 1994; dense population in protected, turfy tundra, Klein Creek, 1747 m, 62°2.29' N 141°19.88' W, C. Roland 95-034, 15 June 1995; scattered in fine, dry reddish soil with gravel, Wiki Creek, 1411 m, 61°54.46’ N 141°10.68’ W, M. Cook 96041, 96099, C. Roland 96-057, 11 June 1996 (ALA); abundant in dry facies around outcrops, Rock Lake, 1119 m, 61°48.7’ N 141°16.57' W, C. Roland 96-038, 6 June 1996; moist, sandy grus with seral herbs on well-vegetated SE-facing alpine colluvium, Horsfeld Creek valley, 1128 m, 62°2’ N 141°11’ W, Parker & Gracz 6913, 13 August 1996. WRANGELL MOUNTAINS: scattered in loose gravel on steep, SE- facing slope, NE slope of Mt. Drum, 1433 m, 62°8.83' N 144°30.18' W, C. Roland 94-051, 11 June 1994. This amphiberingean arctic-alpine fescue is rare in Alaska (G4 $283) where it grows in dry, rocky situations in the subalpine and alpine zones of northern and central Alaska. In the Yukon it is known from numerous stations in the arctic but only a single locality in the southwest corner of the Territory (Cody 1996). The stations cited above extend its range southeast 310 km into the Wrangell Mountains, 320 km into the Mentasta Mountains and 450 km into the Nutzotin Moutains from collections in the Fairbanks Quad (Wood River Buttes, 64°28.35’ N 148°05.97' W, Duffy et al. 95-80, 95-68, 95-26 & 95-57, 16 June 1995 (ALA)). These collections also connect the range to the northwest in Alaska with the range 155 km to the east in the Yukon Territory (Cody 1996). Map 28. Festuca minutiflora Rydb., Small-Flower Fescue — CHUGACH MOUNTAINS: scattered in dry mineral soil of gravel slope, alpine valley near Hanagita Peak, 1186 m, 61°4.91’ N 143°38.86’ W, M. Cook 96418, 7 July 1996 (ALA). This North American cordilleran fescue is new to the flora of Alaska. The main range of this species is in the Rocky Mountains south to California and New Mexico. It is rare in Alaska (G4 $1) and in the Yukon Territory (Douglas et al. 1981). The collection cited above is 445 km west of collections in the Yukon Territory in Kluane National Park (Cody 1996). Map 29. Festuca richardsonii Hooker (F. rubra L. ssp. richardsonii (Hooker) Hultén, Festuca rubra ssp. arctica (Hack.) Govor.), Richardson’s Fescue — THE CANADIAN FIELD-NATURALIST Vol. 116 NUTZOTIN MOUNTAINS: scattered in mesic graminoid stringers between closed low birch willow scrub, Beaver Lake, 1341 m, 62°2.61'’ N 141°48.39’ W, Cook & Roland 94197B, 29 June 1994; scattered on exposed rubble slope, Wiki Creek, 1411 m, 61°54.46' N 141°10.68’ W, C. Roland 96-074, 8 June 1996. WRANGELL MOUNTAINS: gravel bar, Nabesna River, 817 m, 62°18.96’ N 142°54.07’ W, C. Roland 96-195, 20 June 1996; occasional in sand and gravel on river terrace, Jacksina River, 759 m, 62°21.51' N 142°52.87" W.C. Roeland Sa-207 21 June 1996; scattered in Rhacomitrium on gravel bar across from Black Mountain, 945 m, 62°19.02’ N 143°47.44' W, M. Cook 96720, 8 August 1996; sandy area on the edge of alder thicket on floodplain, Mud Lake, 1001.m, 62°13.52".N 143°45: 265 Wo. Roland 96-893, 9 August 1996. This species is circumpolar with an arctic-alpine distri- bution. The specimens cited above connect its range 196 km to the northwest in the Alaska Range (Aiken and Darbyshire 1990) with its range 70 km to the east in the Yukon Territory (Cody 1996). Map 30. Festuca saximontana Rydberg, Rocky Mountain Fescue —- CHUGACH MOUNTAINS: open sandy White Spruce forest, Tana Dunes, 457 m, 61°4.04’ N 142°45.67' W, M. Duffy 91040, 6 July 1991; common on margins of lower Tana Dunes, 488 m, 61°6.19’ N 142°55.03’ W, M. Cook 94199, C. Roland 94-177A, 7 July 1994; occasional clumps in sand of open tall alder scrub, West Fork Tana River, 448 m, 60°52.95' N 142°48.33' W, M. Cook 96565, 7 July 1996; scat- tered among alder-willow patches on moraine, alpine basin of 12-mile Creek, 1271 m, 60°48.85’ N 142°33.52' W, M. Cook 96631, 7 July 1996 (ALA). This fescue occurs across boreal North America and is found as far south as Arizona and New Mexico. Our collec- tions extend its range 338 km south into the Chugach Mountains from a station in the Alaska Range (Hultén 1968) and connect the range 158 km to the east in the Yukon Territory (Cody 1996). Map 31. Glyceria pulchella (Nash) K. Schum., MacKenzie Valley Mannagrass — CHUGACH MOUNTAINS: mar- gin of meadow on river terrace wetlands, Tana River flats, 351 m, 61°12.35’ N 142°51.87' W, Duffy & Barnes 96-188, 8 August 1996. The specimen cited above extends the range of this North American boreal-montane species 337 km south into the Chugach Mountains from the Tanana Valley (Hultén 1968) and connects the range 526 km to the east in the Yukon Territory near Carcross (Cody 1996). Map 32. Phippsia algida (Soland.) R. Br., Snow Grass — CHUGACH MOUNTAINS: scattered in wet gravel, W. Fork Goat Creek, 1487 m, 60°59.8’ N 142°11.8' W, M. Cook 96656, 31 July 1996; moist seepage from snowmelt, Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-298, 29 July 1996 (ALA); scattered in frost polygons, Verde Ridge, 1554 m, 61°14.03’ N 143°28.52’ W, Roland & D’Auria 96- 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 31. Festuca saximontana 32. Glyceria pulchella 215 216 488, 11 July 1996; occasional in moist calcareous gravel, Granite Creek, 1829 m, 61°0.22’ N 141°51.1’ W, C. Roland 96-736, 28 July 1996. WRANGELL MOUNTAINS: scattered in wet silt, Mud Lake, 1001 m, 62°13.52’ N 143°45.28’ W, C. Roland 96-883, 9 August 1996; rare in silt on river bar, Copper River, 847 m, 62°28.75' N 143°38.3’ W, C. Roland 96-934, 12 August 1996. MENTASTA MOUNTAINS: saturated mud seep, headwaters of Lost Creek, 62°36.25’ N 143°12.08’ W, M. Cook 3156, 8 July 1998. NuTzoTIN MounrTAINS: alkali lake bed, Solo Plats, 1335 m, 61°31.55' N 141°25.3" W, M. Duffy 92157, 7 July 1992. ST. ELIAS MOUNTAINS: common in wet seep, 2073 m, 60°57.74’ N 141°17.33' W, M. Cook 95226, C. Roland 95-192, 14 July 1995. WRANGELL MOUNTAINS: rare in moist organic soil, Cone Ridge, 2073 m, 62°8.21’ N 143°18.47' W, M. Cook 94367, C. Roland 94-308, 25 July 1994; occasional in moss, Chetaslina Ridge, 1615 m, 61°56.51' N 144°25.93' W, M. Cook 94470, 15 August 1994; frost scars, VABM Sentinel, 1829 m, 61°39’ N 142°32’ W, Batten & Barker 96- 107, 24 July 1996; moist sand along rivulet, Skolai Creek, 1341 m, 61°41.5’ N 142°23' W, Batten & Barker 96-126, 25 July 1996; common in mossy soil im seep, Masen~ Creek, 1835 my 61°34,06' N 142°18.33' W, C. Roland 96-637, 23 July 1996; locally abundant in saturated soils, Grotto Creek, 1661 m, 61°30.56’ N 142°24.79’ W, C. Roland 96- 657, 23 July 1996; Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5' W, Leggett & Potkin 95-138, 95-108, 28 July 1995; abundant in streambed, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24' W, C. Roland 95-106, 30 June 1995; common in seep, Cooper Mountain, 1942 m, 62°17.16' N 142°31.44’ W, C. Roland 94- 281, 24 July 1994. This circumpolar arctic-alpine tundra grass is rare in the Yukon Territory (Douglas et al. 1981). It had been collect- ed in the Wrangell Mountains at Chitistone Pass (Scott 1968). The collections cited above extend its range 127 km into the Chugach Mountains and 135 km into the western Wrangell Mountains. Map 33. Poa hispidula Vasey (P. macrocalyx Trautv. & C.A. Mey.), Large Glume Blue Grass — CHUGACH MOUNTAINS: common in clumps on boulder strewn area on lower Tana Dunes, 488 m, 61°6.19’ N 142°55.03’ W, M. Cook 94201, 94202, C. Roland 94-177C, 7 July 1994; scattered in saturated, mossy fen, vic. Middle Hanagita Lake, 744 m, 61°11.58’ N 143°30.49' W, Roland & D’Auria 96-498, 12 July 1996. WRANGELL MOUNTAINS: mesic graminoid- dwarf scrub tundra, Black Mountain, 1481 m, 62°20.85' N 143°44.9' W, M. Cook 95160, 3 July 1995. The specimens cited above represent the northern limit of the range of this grass. It is an Alaska-Yukon endemic, known from numerous localities along the southern coast and the Aleutian Islands. These collections extend its range 289 km east into the Chugach Mountains and 264 km THE CANADIAN FIELD-NATURALIST Vol. 116 northeast into the Wrangell Mountains from a collection in the Anchorage Quad (Ft. Richardson, 61°15.45’ N 149°39.42' W, Duffy & Tande 1008, 2 August 1994 (ALA)) and connect the range 406 km to the southeast near Skagway (Hultén 1968). Map 34. Podagrostis aequivalis (Trin.) Scribn. & Merr. (Agrostis aequivalvis (Trin.) Trin.), Northern Bentgrass — WRANGELL MOUNTAINS: rare on flood- plain of the Dadina River, 393 m, 61°51.49’ N 145°0.72’ W, C. Roland 96-308, 1 July 1996. The Wrangell Mountains locality represents the northern limit of the range of this species, which occurs along the west coast of North America, from Oregon to the south coast of Alaska. This locality extends the range of this species 133 km north into the Wrangell Mountains from a station near Cordova (Hultén 1968). Map 35. Puccinellia deschampsioides Sorens., Polar Alkalai Grass — WRANGELL MOUNTAINS: near shoreline in wet clay, Mt. Drum warm spring, 897 m, 62°4.83' N 145°0.55' W, C.R. Meyers 84-90, 18 July 1984; sparsely vegetated berm surrounding Lower Klawasi Mud volcano, 62°3.52' N 145°13.29' W, M. Cook 91203, 91204, 21 August 1991. This halophytic grass is found in disjunct stations across northern North America and western Greenland. The Mt. Drum localities represent its western range limit. These sta- tions are 358 km west of a station in the Yukon Territory (Cody 1996) where it is considered rare (Douglas et al. 1981). Map 36. Puccinellia interior Sorens., Inland Alkalai Grass — NUTZOTIN MOUNTAINS: infrequent in saturated moss and standing water in graminoid dominated marsh adjacent to Ptarmigan Lake airstrip, 1052 m, 61°52.01’ N 141°10.28' W, M. Cook 3198A, 17 July 1998. This species is endemic to Alaska and the Yukon where it occurs in moist meadows. It is known from east and south-central Alaska and the southern half of Yukon Territory, where it is considered rare (Douglas et al. 1981). It was collected in the vicinity of the Park at Copper Center (C.W. Heideman 2, 1908 (US); J.P. Anderson 2734, 1935 (H)) and at Chitina (J.P. Anderson 2028, 7 July 1935 (H)) (Hultén 1941-1950). Our collection connects the distribution from a point 76 km to the east in the Yukon Territory (Cody 1996) with those cited above, approxi- mately 222 km to the west. Map 37. Trisetum sibiricum Rupr. ssp. litorale (Rupr.) Roshev., Siberian Oatgrass — NUTZOTIN MOounrTAINS: S-facing scree slope, 2.4 km west of Ptarmigan Creek, 1494 m, 61°32.89’ N 141°3.28' W, M. Cook 92482, 9 August 1992; steep S-facing rhyo- lite scree knob, west side of Wiki Creek, 1524 m, 61°54.77' N 141°11.05’ W, M. Cook 3190, 15 July 1998 (ALA). This amphiberingean species occurs throughout northern Asia, but in North America is restricted to Alaska and a few sites in westernmost Yukon Territory where it is considered very rare (Douglas et al. 1981). This grass is also consid- ered rare in Alaska (GST4Q $2). Our collections extend its range 515 km south into the Nutzotin Mountains and repre- sent the southern limit of its distribution in North America. Map 38. 217 NOTABLE VASCULAR PLANTS FROM ALASKA CooK AND ROLAND 2002 da Phippsia algi 53. idula Poa hisp 34. ivalis Podagrostis aequ a3. Puccinellia deschampsioides 36. 218 Vahlodea atropurpurea (Wahlenb.) E. Fries, Mountain Hairgrass — GULF OF ALASKA: mesic tall mixed forb herbaceous meadow, Amphitheater Knob, 427 m, 59°57’ N 139°46' W, M. Cook 87-88, 19 August 1987; mixed herbaceous meadow, Karr Hills, 549 m, 60°8.66’ N 141°16.44' W, K. Beck 87- 7, 12 August 1987; heather alpine tundra, Floral Pass, 792 m, 59°57.91’ N 139°57.76’ W, M. Cook 87-100, 21 August 1987; Samovar Hills, 518 m, 60°9.3’ N 140°38’ W, M. Cook 87-105, 23 August 1987. CHUGACH MOUNTAINS: moist soil in Betula glandulosa thicket, Martin Creek, 60°56’ N 142°23’ W, Batten & Barker 96-260, 28 July 1996; scattered in tall forb herbaceous meadow, Granite Creek, 1344 m, 60°46.44’ n 142°5.12' W, C. Roland 96-805b, 96-804b, M. Cook 96224, 29 July 1996; common in Empetrum heath, 12-Mile Creek, 988 m, 60°50.51' N 142°23.02’ W, C. Roland 96-764, 29 July 1996; common in subarctic lowland sedge wet meadow, upper Tebay Lake, 579 m, 61°11’ N 144°24’ W, Parker & Gracz 6769, 6778, 7 August 1996. WRANGELL MOUNTAINS: patchy in mesic low willow forb herbaceous vegetation, Lakina Glacier, 1219 m, 61°33.64'’ N 143°19.01’ W, M. Cook 96540, C. Roland 96-700, 25 July 1996; patchy along shore of lake in Empetrum heath, Cheshnina Plateau, 1399 m, 61°48.04’ N 144°6.27' W, M. Cook 96717, 8 August 1996. St. ELIAS MOUNTAINS: open low shrub, Short River Pond, 503 m, 61°5.35’ N 141°56.3' W, Parker & Duffy 6695, 6 August 1996. Vahlodea atropurpurea sens. lat. occurs in meadows and thickets throughout the circumpolar north with large gaps (Cody 1996). The specimens cited above extend its range 401 km to the south into the southern St. Elias Mountains from a station near Slana (Hultén 1968) and connect the western range with a station 385 km to the southeast in the Sitka Quad (Freshwater Creek, 58°0.06’ N 134°49.57' W, C.L. Parker 5251, 14 July 1994 (ALA)). Map 39. CYPERACEAE Carex adelostoma Krecz., Circumpolar Sedge — WRANGELL MOUuNTAINS: Fox Farm Lakes, 727 m, 62°19.98’ N 144°50.02’ W, M. Potkin 95-008, 26 July 1995; patchy in mesic graminoid herbaceous vegetation, vic. Copper Lake, 911 m, 62°24.53’ N 143°42.68' W, M. Cook 96732, 8 August 1996. Carex adelostoma was collected by A. Dutilly, E. Lepage and O’Neil near Long Lake on the Nabesna Road in 1947 (Lepage 1951; Hultén 1967). Our two collection localities are nearby in the upper Copper River watershed. These three collections represent the only known localities in Alaska and form the western range limit of this species in North America. This sedge is rare in Alaska (G4 S1) and has an interrupted distribution throughout its range. The closest station is 988 km to the northeast at Great Bear Lake in the Northwest Territories (Porsild and Cody 1980). . Map 40. Carex albo-nigra Mack., Black-and-White-Scale Sedge — CHUGACH MOUNTAINS: S-facing boulder slope in unstable gravel scree, Juniper Island, 1291 m, 60°36.24’ N 142°21.69' W, M. Cook 94264, THE CANADIAN FIELD-NATURALIST Vol. 116 11 July 1994. MENTASTA MOUNTAINS: scattered in Arctostaphylos uva-ursi/Festuca altaica meadow on S-facing slope, 1097 m, 62°33.3’ N 143°9’ W, Roland & D’Auria 97-042, 26 June 1997; common in dry, White Spruce forest at base of bluff, Lost Creek, 1097 m, 62°33.3’ N 143°9’ W, Roland & D’Auria 97-077, 26 June 1997. NUTZOTIN MOUNTAINS: occasional on lower slopes in Potentilla fruticosa/Arctostaphylos uva-ursi community, Carl Creek, 1920 m, 62°3.52' N 141°36.27' W, C. Roland 95-043, 20 June 1994; crevices of boulder outcrops on S-facing slope, ridge 0.6 km E of Rocker & Ptarmigan Creek confluence, 1433 m, 61°54.8’ N 141°2.03' W, M. Cook 94142A, 23 June 1994; patchy in talus meadow of S-facing slope, Rock Lake, 1119 m, 61°48.7' N 14]°0Ree C. Roland 96-034, 5 June 1996. WRANGELL MOUNTAINS: wet graminoid-dwarf willow-bryoid tundra, ridge between Chichokna and Chetaslina Rivers, 1615 m, 61°56.51’ N 144°25.93’ W, M. Cook 94470A, 15 August 1994. This sedge occurs on dry hillsides and talus meadows into the subalpine region in eastern Alaska. It is endemic to the cordilleran region of western North America, where it occurs from New Mexico, Arizona and California north- ward into eastern Alaska. It is considered rare in the Yukon Territory where it is known from fewer than ten sites (Douglas et al. 1981). Our collections extend its range 299 km southeast into the Nutzotin Mountains and 365 km into the Chugach Mountains from a locality in the Alaska Range (Black Rapids Roadhouse, 63°32.0’ N 145°51.0’ W, Batten et al. 78-4, 28 June 1978 (ALA)). Map 41. Carex buxbaumii Wahlenb., Buxbaum’s Sedge — CHUGACH MOUNTAINS: sedge meadow, Upper Tebay Lake, 579 m, 61°11.5' N 144°23" W, 2. Ab Viereck 11093, 15 July 1996; rare in sedge herb meadow, Tebay Lake, 579 m, 61°11’ N 144°24’ W, Parker & Gracz 6740, 7 August 1996. NUTZOTIN MOUNTAINS: patchy in sphagnum bog, Lick Creek, 914 m, 62°28.89’ N 142°7.56’ W, M. Cook 95111, 28 June 1995. WRANGELL MOUNTAINS: patchy in mesic graminoid herbaceous-open low willow scrub gulley, vic. of Whitham Lake, 936 m, 62°19.33' N 142°53.28’ W, M. Cook 96200, 6 June 1996; abun- dant at edge of pond, Lakina river, 792 m, 61°29.49’ N 143°25.6' W, C. Roland 96-678, 24 July 1996; occasional in marsh, Dadina River, 744 m, 61°53.05’ N 144°44.09' W, M. Cook 96276A, C. Roland 96- 344, 2 July 1996. This sedge is found in swamps and bogs at low eleva- tions across boreal regions of the circumpolar north. However, it is absent from the Russian Far-East according to Hultén (1968). In North America, it occurs as far south as Georgia in the east and California in the west. It is of limited distribution in the Yukon Territory (Cody 1996). The localities cited above extend its range 189 km to the south into the Chugach Mountains from a station near Mentasta reported by Hultén (1968). Map 42. Carex chordorrhiza Ehrh. ex. L. f., Creeping Sedge — CHUGACH MOUNTAINS: sweetgale/graminoid bog, 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 219 40. Carex adelostoma 220 Tana River flats, 411 m, 61°12.59’ N 142°54.47’ W, Duffy & Barnes 96-194, 8 August 1996. WRANGELL MOunrTAINS: numerous in quaking bog, Indian Lake, 655 m, 62°38.1' N 144°20.2' W, M. Cook 95347, 1 August 1995; locally dominant in wet sedge mead- ow, vic. Monte Cristo Creek, 1067 m, 62°14.27' N 142°55.81' W, C. Roland 96-136, 18 June 1996; co-dominant in sedge marsh, Kuskulana River, 509 mi, 61°28.53', N L43"50772 WC. Roland 96-722, 26 July 1996. This sedge occurs in bogs and lake margins from the lowlands into the subalpine region in Alaska. Its range is circumpolar, but in North Ameica it is generally restricted to the boreal region and northward, reaching its southern range limit in isolated localities in Illinois. Our collections form a range connection for this sedge between a station in the Yukon Territory 255 km to the east (Cody 1996) and a locality in the Anchorage Quad 226 km to the west (Jim and Swan Lakes, 61°33.0' N 148°55.0’ W, Batten & Reed 80-115, 1 July 1980 (ALA)). Map 43. Carex crawfordii Fern., Crawford’s Sedge — CHUGACH MOUNTAINS: dry pond meadow, Tana River flats, 442 m, 61°14.45’ N 142°57.21' W, Duffy & Barnes 96-140, 8 August 1996. This sedge occurs across boreal North America and reaches its southern range limit in Missouri (Kartesz 1999). It is rare in Alaska (G5 $2S3) and in the Yukon Territory (Douglas et al. 1981). The closest collection localities to the Tana River are a station near Delta Junction (141 km north) and a station near Anchorage 331 km to the west (Hultén 1968). Map 44. Carex eburnea Boott, Bristleleaf Sedge — ST. ELIAS MOUunNrTAINS: White Spruce woodland, Clear Stream, 488 m, 61°5.82’ N 141°57.92’ W, Duffy & Barnes 96-001, 8 August 1996. WRANGELL MOUNTAINS: patchy in sedge meadow, Grizzly Lake, 1000 m, 62°19.51' N 143°9.24' W, Cook & Allen 3551, 26 July 2000. These collections extend the range of this North American boreal-montane sedge 182 km south into the Wrangell Mountains from the Alaska Range (Hultén 1968). It had been collected near Chitina (Hultén 1968) and at the head of the Chitina River (H.M. Laing s.n., 18 June 1925 (CAN), Hultén 1941-1950, Porsild 1939). It is rare in Alaska (G5 $2S3) and in the Yukon Territory (Douglas et al. 1981). Map 45. Carex filifolia Nutt., Thread-Leaf sedge — CHUGACH MOUNTAINS: steep, SE-facing bluff, confluence of Copper and Chitina Rivers, 152 m, 61°31.55’ N 144°24.85' W, C. Roland 94-074B, 12 June 1994; steep SW-facing scree slope, Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-187B, 96-224, 27 & 28 July 1996. MENTASTA MOUNTAINS: rare in Arctostaphylos uva-ursi mats on S-facing slope, Devil’s Mountain, 942 m, 62°24.95' N 142°54.86'’ W, C. Roland 96-231, 22 June 1996. NUTZOTIN MOUNTAINS: abundant on steep, S-facing bluff, White River between Cub and Traver Creeks, 1219 m, 61°44.3’ N 141°9.5' W, C. Roland 95-078, 21 June 1995; occasional in well drained areas on THE CANADIAN FIELD-NATURALIST Vol. 116 limestone, Mt. Natazhat, 1716 m, 61°35.38' N 141°1.83’ W, C. Roland 95-060, 19 June 1995. ST. ELIAS MOUNTAINS: dry sandy mineral soil, vic. Walsh Glacier, 1951 m, 60°53.94’ N 141°6.75’ W, M. Cook 95222, 15 July 1995. Carex filifolia occurs throughout the western half of North America, reaching its eastern limit in Manitoba, Canada and its southern limit in New Mexico. It occurs on xeric slopes and prairies, and only reaches into eastern Alaska, where Hultén reported it from two localities, one near Chitina (D.F. Murray, B.A. Murray and A.P. Khokhryakov s.n., 61°30.0' N 144°25.0’ W, 8 July 1981 (ALA)) and another in the Tanana River valley east of Fairbanks. Our collections extend its range into the Nutzotin, Chugach, St. Elias and Wrangell Mountains. Map 46. Carex holostoma Drej., Arctic Marsh Sedge — NUTZOTIN MOUNTAINS: occasional in open low mixed shrub sedge tundra bog, Horsfeld Creek, 1097 m, 62°1’ N 141°11’ W, Parker & Gracz 6857A, 10 August 1996. The specimen cited above extends the range of this sedge 134 km east into the Nutzotin Mountains from a col- lection made by A. Dutilly, E. Lepage and O’Neil near Long Lake on the Nabesna Road in 1947 (Lepage 1951; Hultén 1967). These collections document the southern limit of its range, which is 833 km south of collections on the arctic coast of Alaska (Mt. Michelson Quad, Kavik R.., 69°41.0' N 146°52.0' W, M. Emers s.n., 2 July 1992 (ALA)). This sedge is rare in Alaska (G3G4 S2) and in the Yukon Territory (Douglas et al. 1981). Map 47. Carex interior Bailey, Inland Sedge — CHUGACH MOUNTAINS: occasional in open mossy areas, West Fork Tana River, 448 m, 60°52.95’ N 142°48.33’ W, M. Cook 96567, 7 July 1996 (ALA). This sedge is disjunct from the temperate zone and is rare in Alaska (GS S1) and in the Yukon Territory (Douglas et al. 1981). The specimen cited above extends its range 288 km east into the Chugach Mountains from a station in the Anchorage Quad (Baxter Bog, 61°13.0’ N 149°54.0’ W, E. F. Layser 3275, July 1984 (ALA)). Map 48. Carex krausei Boeck., Krause’s Sedge — CHUGACH MOUNTAINS: wet sedge meadow on low terrace, Kiagna River, 762 m, 60°58.33’ N 142°20' W, Batten & Barker 96-239, 28 July 1996. MENTASTA MOunraAINs: landslide on W end of Soda Lake, 1173 m, 62°32.36' N 142°53.98’ W, M. Cook 94306B, 21 July 1994; occasional in wet sand in openings of Populus trichocarpa/Salix alaxensis woodland, Totschunda Creek, 725 m, 62°26.94’ N 42°40.8’ W, M. Cook 96254, 6 June 1996. NUTZOTIN MOUNTAINS: scattered under Salix pulchra, upper Flat Creek, 1323 m, 61°58.97' N 141°41.25’ W, M. Cook 95036, 17 June 1995; St. ELIAS MOUNTAINS: open low wil- low scrub, Clear Stream, 488 m, 61°5.82'’ N 141°57.92' W, Duffy & Barnes 96-029, 96-030, 96- 031, 8 August 1996. WRANGELL MOUNTAINS: few in bare gravel patches, vic. Boulder Creek, 1036 m, 62°31.27' N 144°11.26’ W, M. Cook 95359, 2 August 1995; occasional in moist silt of abandoned river channel, vic. Black Mountain, 945 m, 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 24 41. Carex albo-nigra 44. Carex crawfordii 222 THE CANADIAN FIELD-NATURALIST Voli a6 47. Carex holostoma 48. Carex interior 225 NOTABLE VASCULAR PLANTS FROM ALASKA CooK AND ROLAND 2002 1 Carex krause 49. . Americana locarpa Ssp Carex las 50. ee Carex laxa a1; Carex lenticularis var. dolia 52. 224 62°19.02' N 143°47.44’ W, C. Roland 96-901, 10 August 1996; patchy in mudflats between gravel bar and spruce woodland, Copper River at outflow creek from Lake 2990, 884 m, 62°24.38' N 143°42.28’ W, M. Cook 96724, 8 August 1996. This circumpolar arctic-alpine sedge had been collected at the head of the Chitna River (H.M. Laing 21, 1925 (CAN) (Hultén 1941-1950; Porsild 1939)). The collections cited above extend its range into the Wrangell, Mentasta and Nutzotin Mountains and connect the distribution 243 km to the north in the Alaska Range (Caster Glacier, 63°24.0’ N 145°43.0'’ W, Palmer & Porsild 467, 27 June 1926 (ALA)). Map 49. Carex lasiocarpa Ehrh. ssp. americana (Fern.) Hult., Wooly-Fruit Sedge — CHUGACH MOUNTAINS: river terraced wetlands surrounded by White Spruce for- est, Tana River flats, 442 m, 61°14.45’ N 142°57.21' W, Duffy & Barnes 96-118, Parker & Gracz 6788, 8 August 1996. WRANGELL MOUNTAINS: dense swards around small pond, Chokosna Lake; 619 m, 61°27.45'’ N 143°49' W, C. Roland 94-330, 10 August 1994; abundant in wet sedge meadow, vic. Dadina River, 744 m, 61°53.05’ N 144°44.09’ W, C. Roland 96-339, 2 July 1996; dominant in pond, Kuskulana River, 509 m, 61°28.53’ N 143°50.77' W, C. Roland 96-720, 96-723, 26 July 1996. This North American boreal-montane species is rare in the Yukon Territory (Douglas et al. 1981). The specimens cited above extend its range east 231 km into the Copper and Chitina River basins from a collection in the Anchorage Quad (Otter Lake, 61°17.53' N 149°44.17' W, Duffy & Tande 1017, 3 Aug 1994 (ALA)). Map 50. Carex laxa Wahlenb., Weak Sedge — NUTZOTIN MOUNTAINS: scattered in marsh, Lick Creek, 914 m, 62°28.89' N 142°7.56' W, Cook & Beck 95112A, 29 June 1995. WRANGELL MOUNTAINS: graminoid meadow, confluence of Monte Cristo Creek and Nabesna River, 1067 m, 62°14.27' N 142°55.81' W, C. Roland 96-134, 18 June 1996. The collections of this boreal-montane sedge cited above are the only verified stations in Alaska. David F. Murray reviewed the collection from Mile 172-174 on the Richardson Highway (J. P. Anderson 2712B (S), Hultén 1941-1950) but could not confirm the original determina- tion from the material (personal communication with D. F. Murray, 11 July, 2000). The collections cited above extend its range 368 km to the west from the Yukon Territory. This sedge is rare in Alaska (G4 S1) and in the Yukon Territory (Douglas et al. 1981). Map 51. Carex lenticularis Michx. var. dolia (M.E. Jones) L.A. Standley. comb. nov. (C. enanderi Hult.), Lakeshore Sedge — CHUGACH MOUNTAINS: flooded silt bar of tributary stream, Martin Creek, 1097 m, 60°56’ N 142°23' W, Batten & Barker 96-266, 28 July 1996; scattered in snowmelt rivulet, west fork - of 12-mile Creek, 1326 m, 60°50.21'’ N 142°30.85’ W, C. Roland 96-787, 29 July 1996; marsh on allu- vial fan, Granite Creek, 823 m, 60°44’ N 142°13’ W, Batten & Barker 96-343, 30 July 1996; moist mead- ow, west fork of 12-mile Creek, 1326 m, 60°50.21’ THE CANADIAN FIELD-NATURALIST Vol. 116 N 142°30.85’ W, M. Cook 96593, 7 July 1996 (ALA). This North American cordilleran sedge is rare in Alaska (G5T3Q S3). The specimens cited above extend its range 188 km to the east of a collection in the Valdez Quad (Thompson Pass, 61°08.0' N 145°45.0’ W, C.L. Parker 2451, 23 July 1990 (ALA)) and connect the range 423 km to the southeast near Skagway (Hultén 1968). Map 52. Carex leptalea Wahlenb., Bristly-stalked Sedge — CHUGACH MOUNTAINS: sweetgale-graminoid bog, Tana River, 411 m, 61°12.59’ N 142°54.47' W, Duffy & Barnes 96-196, 8 August 1996; scattered to common in small patch of wet sedge herb tundra, Tana River, 335 m, 61°11’ N 142°51’ W, Parker & Gracz 6820, 9 August 1996; common in elevated mossy hummocks in Sphagnum bog, Hanagita River, 762 m, 61°11.07’ N 143°25.81' W, Roland & D’Auria 96-493A, 12 July 1996; common in elevat- ed, somewhat drier margin of wet graminoid mead- ow, Nerelna Creek, 884 m, 61°25.3’ N 144°18.61’ W, Roland & D’Auria 96-424, 9 July 1996. GULF OF ALASKA BASIN: horsetail bog, Manby Beach, 59°47.52’ N 140°56.51’ W, M. Cook 87-117, 20 August 1987. NUTZOTIN MOUNTAINS: scattered in marsh, Lick Creek, 914 m, 62°28.89’ N 142°7.56’ W, M. Cook 95113, 29 June 1995. WRANGELL MOUNTAINS: clumped in Sphagnum at margin of pond, Lakina River, 792 m, 61°29.49' N 143°25.6' W, M. Cook 96522, 7 July 1996; elevated hummocks in wet sedge meadow, Dadina River, 744 m, 61°53.05' N 144°44.09’ W, C. Roland 96-341, 2 July 1996. This sedge is North American with a boreal-montane dis- tribution. The localities cited above extend its range south 220 km into the Nutzotin Mountains, 266 km into the Wrangell Mountains and 296 km into the Chugach Mountains from a station in the Alaska Range (Hultén 1968). The collection from the Malaspina Forelands extends the range 433 east of a station on the Kenai Peninsual (Hultén 1968). Map 53. Carex nardina E. Fries, Nard Sedge — CHUGACH MOUNTAINS: rock outcrops, Chakina River, 1646 m, 61°5.31’ N 143°0.53’ W, M. Cook 94213, 8 July 1994; common in Dryas-sedge tundra, Juniper Island, 1291 m, 60°36.24' N 142°21.69’ W, M. Cook 94271, 12 July 1994; SW-facing rock crevice, Juniper Island, 1346 m, 60°36.17’ N 142°14.96' W, Cook & Roland 94-205, 94250A, 10 July 1994; occa- sional in marble rubble, Canyon Creek, 1463 m, 61°24.35' N 144°20.8' W, Roland & D’Auria 96- 437, 9 July 1996. MENTASTA MOUNTAINS: scattered in tundra, Totschunda Creek, 1280 m, 62°27.63'’ N 142°12.44' W, C. Roland 96-277, 24 June 1996; dry tundra on S-facing slope, Devils’s Mountain, 1530 m, 62°25.62’ N 142°53.93’ W, C. Roland 96- 360C, 3 July 1996; common on limestone outcrops, Trail Creek, 1615 m, 62°36.05’ N 143°17.63’ W, C. Roland 95-005, 6 June 1995. NUTZOTIN MOUNTAINS: limestone gravel, Baultoff Creek, 1707 m, 62°9.13' 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 56. Carex obtusata 225 226 N 141°14.51’ W, C. Roland 94-164, 28 June 1994; limestone ridge, Lime Butte, 1554 m, 61°21.56’ N 142°26.43' W, Cook & Roland 94-232B, 94298B, 14 July 1994. St. EL1as MOUNTAINS: occasional on limestone gravel, Mt. Natazhat, 1716 m, 61°35.38’ N 141°1.83' W, M. Cook 95069, 19 June 1995; com- mon in disturbed marble rubble, Blondie Ridge, 1951 m, 60°53.94’ N 141°6.75' W, C. Roland 95- 152, 11 July 1995. WRANGELL MOUNTAINS: gravelly limestone, Chitistone Mountain, 1219 m, 61°28’ N 142°33’ W, Batten & Barker 96-141, 26 July 1996; scattered in gravelly tundra, Cheshnina Plateau, 1399 m, 61°48.04’ N 144°6.27' W, C. Roland 96-854, 6 August 1996. This sedge is amphi-atlantic with an arctic-alpine distri- bution. It was collected in the Wrangell Mountains at Bonanza Ridge (Nordell and Schmitt 1978) and Guerin Glacier (61°37.0' N 141°50.0’ W, D.F. Murray 2066, 3 August 1968 (ALA)). The localities cited above extend its range 118 km east from a station in the Alaska Range (Hultén 1968) and 102 km south into the Chugach Mountains from the Bonanza Ridge collection. Map 54. Carex nigricans C.A. Mey, Blackish sedge — CHUGACH MOUNTAINS: alpine herbaceous meadow, Karr Hills, 549 m, 60°8.66’ N 141°16.44’ W, K.A. Beck 87-11, 12 August 1987; occasional in Empetrum heath, 12-mile Creek, 988 m, 60°50.51' N 142°23.02' W, C. Roland 96-766A, 29 July 1996; scattered in snowmelt rivulet, 12-mile Creek, 1326 m, 60°50.21' N 142°30.85' W, C. Roland 96- 788, 29 July 1996; scattered in lush forb-herbaceous meadow, Granite River, 1344 m, 60°46.44’ N 142°5.12' W, C. Roland 96-800, 29 July 1996; occa- sional in wet moss, vic. Spirit Mountain, 1006 m, 61°16.84’ N 144°29.86' W, M. Cook 96433, 7 July 1996; occasional in Cassiope/Luetkea heath of alpine basin, vic. 12-mile Creek, 1271 m, 60°48.85' N 142°33.52' W, M. Cook 96626, 7 July 1996. GULF OF ALASKA: Cassiope alpine tundra, Samovar Hills, 549 m, 60°8.11’ N 140°39.55’ W, M. Cook 87-102, 23 August 1987; gravelly area next to wet equisetum meadow, Guyot Glacier, 183 m, 60°3.6’ N 141°18’ W, K.A. Beck 92545, 22 August 1992. WRANGELL MOUNTAINS: ridge 0.6 km S of Nadina Glacier, 1768 m, 62°2.85' N 144°41’ W, M. Cook 94424A, 30 August 1994. The northern limit of the range of this North American cordilleran sedge is documented by the collection cited above from the Wrangell Mountains. These collections extend its range 116 km to the northeast into the Wrangell Mountains, 203 km east into the Chugach Mountains and 291 km southeast into the southern St. Elias Mountains from a station near Thompson Pass (Hultén 1968) and con- nect the range 298 km to the southeast near Skagway (Hultén 1968). Map 55. Carex obtusata Liljeb., Blunt Sedge — CHUGACH MOUNTAINS: rubbly area of SW-facing slope underlain by ultramafic rock, Chakina River, 1768 m, 61°6.19’ N 142°55.03' W, C. Roland 94-185B, 8 July 1994; dry THE CANADIAN FIELD-NATURALIST Vol. 116 slopes, Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-213, 27 July 1996; dry soil on exposed bench, Goat Creek, 1487 m, 60°59.5’ N 142°1' W, Batten & Barker 96-316, 29 July 1996 (ALA). NUTZOTIN MOUNTAINS: scattered in turfy, Kobresia dominated community on knoll, Rock Lake, 1119 m, 61°48.7' N 141°16.57' W, C. Roland 96-033, 5 June 1996. WRANGELL MOUNTAINS: Dryas/dwarf willow tundra community, Moose Point, 1052 m, 62°31.61' N 144°12.38' W, C. Roland 94-021, 7 June 1994; occasional on dry hillside, Nabesna River bluff, 1106 m, 62°15.05’ N 142°54.68’ W, C. Roland 96-157, 19 June 1996; few in bare mineral soil on well drained slope, Nabesna River bluff, 1106 m, 62°15.05' N 142°54.68’ W, M. Cook 96192, 6 June 1996. This species is amphiberingean with an arctic-alpine distribution. The specimens cited above extend its range south 169 km into the Wrangell Mountains, 244 km into the Nutoztin Mountains and 315 km into the Chugach Mountains from a collection in the Alaska Range (Mt. Hayes Quad: 63°43.0' N 144°23.0’ W, Batten & Juday 85-207, 12 July 1985 (ALA)) and connect the range 126 km to the east in the Yukon Territory (Cody 1996). Map 56. Carex parryana Dew., Parry’s Sedge — ST. ELIAS MOUNTAINS: open low willow scrub, Clear Stream, terminus of Barnard Glacier, 488 m, 61°5.82’ N 141°57.92' W, Duffy & Barnes 96-012, 8 August 1996. This North American boreal-montane sedge is rare in Alaska (G4 S1). The specimen cited above extends the range in the Chugach Mountains 348 km to the east from a collection in the Anchorage Quad (Eklutna, 61°28.0' N 149°22.0' W, Dutilly et al. 20709, 7 July 1947 (ALA)) and is 162 km west of collections in the Yukon Territory (Cody 1996). Map 57. Carex pauciflora Lightf., Star Sedge — CHUGACH MOUNTAINS: boggy area in open tall willow alder scrub, Falls Creek, 610 m, 61°10.58'’ N 144°24.75’ W, M. Duffy 91103, 12 August 1991; ericaceous shrub bog, Middle Fork of the Bremner River, 869 m, 60°55.05’ N 143°43.86’ W, Duffy & Barnes 96-093, 8 August 1996. GULF OF ALASKA: ericaceous shrub/graminoid herbaceous bog, Robinson Mountains, 427 m, 60°4.8’ N 142°12.8' W, M. Duffy 9216, 11 June 1992. The collections cited above extend the range of this bore- al-montane sedge east 111 km in the Chugach Mountains and 206 km into the Robinson Mountains from a station near Cordova (Orca, 60°39.83’ N 145°43.00’ W, LL. Norberg s.n., 28 August 1938 (S), Hultén 1941) and connect the range 227 km to the southeast near Yakutat (Lost River area, 59°28.0’ N 139°40.0' W, C. L. Parker 2611, 24 June 1991 (ALA)). Map 58. Carex petasata Dew., Liddon Sedge —- CHUGACH MOUNTAINS: locally common in a dry meadow on lateral moraine, Granite Creek, 884 m, 60°44.63’ N 142°6.05' W, C. Roland 96-791, 29 July 1996 (ALA). 2002. COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 227 60. Carex petricosa 228 This North American cordilleran sedge is new to the flora of Alaska and considered rare (G5 S1). The collection cited above is 262 km west of stations in the southwest Yukon near Haines Junction (Cody 1996) and represents the western limit of its known range. Map 59. Carex petricosa Dewey, Rockdwelling Sedge — CHUGACH MOUNTAINS: dry soil on steep SW-facing slope, Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-219, 27 July 1996; occasional patches in gravel and finer scree, vic. Iron Creek, 1612 m, 61°1.49’ N 141°54.23’ W, M. Cook 96580, 7 July 1996. MENTASTA MOUNTAINS: densely vegetated shoulder amid extensive gravel chutes on SW-facing slope, Totschunda Creek, 1280 m, 62°27.63’ N 142°12.44’ W, C. Roland 96-296, 24 June 1996. NUTZOTIN MOUNTAINS: subalpine meadow at treeline, Carden Hills, 1311 m, 62°18.44’ N 141°11.55’ W, C. Roland 94-135, 25 June 1994. St. ELIAS MOUNTAINS: common in dry tundra on limestone, ridge between Dan. & Copper Creeks, 1554 m,-61°21:56' N 142°26.43' W, C. Roland 94-231B, 14 July 1994; occasional in well-drained areas on limestone, Mt. Nataziiat, 1716 m; G1°35-38"- NM 1183" W. Cc. Roland 95-059, 95-061, 19 June 1995; in limestone scree with stringers of Dryas alaskensis, Mt. Natazhat, 1716 m, 61°35.38’ N 141°1.83’ W, M. Cook 95078A, 21 June 1995. WRANGELL MOUNTAINS: occa- sional in open, subalpine White Spruce forest and meadows on W-facing slope of limestone knoll, Fish Creek, 1067 m, 62°16.92' N 142°58.99' W, C. Roland 95-305, 30 July 1995. This North American cordilleran sedge had been collect- ed in the St. Elias Mountains (Guerin Glacier, 61°37.42’ N 141°4.38' W, D. F. Murray 2063, 3 August 1968 (ALA)). The collections cited above extend its range 91 km south into the Chugach Mountains, 111 km north into the Nutzotin Mountains and 126 km northwest into the Wrangell Mountains. These collections also connect the range 203 km to the northwest in the Alaska Range (Mile 238 Richardson Highway, 63°40.0’ N 145°52.0' W, Batten & Dawe 78-123, 4 July 1978 (ALA)) with the range 134 km to the east in the Yukon Territory (Cody 1996). Map 60. Carex phaeocephala Piper, Dunhead Sedge — CHUGACH MOUNTAINS: scattered in loose, shaly talus on S-facing slope, vic. 12-mile Creek, 1335 m, 60°49.71' N 142°33.34’ W, C. Roland 96-815, 30 July 1996; tufted on slope of moraine in Arctostaphylos uva-ursi patches, Granite River, 884 m, 60°44.63’ N 142°6.05’ W, M. Cook 96602, 7 July 1996; mesic shrub birch-ericaceous, Granite Creek, near Ross Green Lake, 701 m, 60°43.83’ N 142°28.78' W, Duffy & Barnes 96-056, 96-057, 8 August 1996. This North American cordilleran sedge is rare in the Yukon Territory (Douglas et al. 1981) and known from only five localities in Alaska. The collections cited above extend its range 104 km east in the Chugach Moutains from a collection near Cordova (Schwan Glacier, 60°58.0’ N 145°00.0' W, C. Parker 1763, 12 August 1986 (ALA)) and THE CANADIAN FIELD-NATURALIST Vol. 116 connect the range 134 km to the east in the Yukon Territory (Cody 1996). Map 61. Carex praticola Rydb., Meadow Sedge — CHUGACH MOUNTAINS: silt of ledges, Juniper Island, 1346 m, 60°36.17' N 142°14.96’ W, Cook & Roland 94-204, 94254, 10 July 1994; scree on S-facing boulder slope, Juniper Island, 1291 m, 60°36.24’ N 142°21.69’ W, M. Cook 94260, 11 July 1994; moist gravel, Short River Pond, 503 m, 61°5.35’ N 141°56.3’ W, Parker & Duffy 6686, 6 August 1996; edges of pools in marshy area, Granite River, 823 m, 60°44’ N 142°13’ W, Batten & Barker 96-344, 30 July 1996. This sedge is North American with a boreal-montane distribution. The localites cited above extend its range 244 km east in the Chugach Mountains from a station near Valdez (Hultén 1968) and connect its range 283 km to the east in the Yukon Territory near Haines Junction (Cody 1996). Map 62. Carex stylosa C.A. Mey, Variegated Sedge — CHUGACH MOUNTAINS: muddy margin of small kettle pond, Granite Creek, 701 m, 60°43.78’ N 142°31.4’ W, Duffy & Barnes 96-066, 8 August 1996; scattered in wet meadow, Granite River, 884 m, 60°44.63’ N 142°6.05' W, C. Roland 96-795B, 29 July 1996; sedge meadow, Upper Tebay Lake, 579 m, 61°11.5’ N 144°23’ W, L. A. & E. G. Viereck 11096, 7 July 1996; mesic sedge herb meadow, Tebay Lake, 579 m, 61°11’ N 144°24’ W, Parker & Gracz 6725, 6767A, 7 August 1996; locally common in mossy heath, 12-Mile Basin, 1335 m, 60°49.71’ N 142°33.34' W, C. Roland 96-830, 30 July 1996; eri- caceous shrub bog, Middle Fork of the Bremner River, 869 m, 60°55.05’ N 143°43.86' W, Duffy & Barnes 96-091, 96-100, 8 August 1996. WRANGELL MOUNTAINS: common in mud on tundra pond, Cheshnina Plateau, 1399 m, 61°48.04’ N 144°6.27' W, C. Roland 96-857, 6 August 1996. This amphiberingean boreal-montane sedge is rare in the Yukon Territory (Douglas et al. 1981). The collections cited above extend its range 135 northwest into the Wrangell Mountains and 214 east into the Chugach Mountains from a station near Worthington Glacier (61°1.08’ N 146°34’ W, L.A. Viereck 8475, 1 August 1967 (ALA)). Map 63. Carex viridula Michx. (C. oederi Retz. ssp. viridula (Mischx.) Hult.), Little Green Sedge — CHUGACH MOUNTAINS: rare in wet silt on floodplain, conflu- ence of Copper and Bremner Rivers, 152 m, 60°56.95' N 144°41.79’ W, C. Roland 96-610, 17 July 1996. GULF OF ALASKA: wet sedge bog, Sudden Stream, 30 m, 59°30.71' N 139°46.44’ W, M. Cook 8850, 20 July 1992; gravelly area next to wet Equisetum meadow, Guyot Glacier, 183 m, 60°3.6’ N 141°18’ W, M. Cook 92552, 19 August 1992. St. ELIAS MOUNTAINS: wet fen in black spruce woodland, Bryson Bar, 1800 m, 61°4.57' N 141°54.47’ W, M. Duffy 91029, 3 July 1991; mixed 2002. COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 61. Carex phaeocephala ASR X \ BON So oo as vague See o “ cee 63. Carex stylosa 64. Carex viridula 229 230 graminoid-forb meadow, Clear Stream, 488 m, 61°5.82’ N 141°57.92’ W, Duffy & Barnes 96-034, 8 August 1996. WRANGELL MOUNTAINS: mud flats around Lake 2870, vic. Tanada Creek, 875 m, 62°31.38' N 143°28.12' W, Moran & Roland 95-08, 28 June 1995. This amphiberingean boreal-montane sedge is rare in the Yukon Territory (Douglas et al. 1981) and known from few localities in Alaska. The collections cited above extend its range 190 km south into the Wrangell Mountains from a sta- tion in the Alaska Range (Hultén 1968), 337 km east in the Chugach Range from a collection in the Anchorage Quad (Bulldog Road bogs, 61°13.10' N 149°41.40’ W, Duffy & Tande 1065, 5 August 1994 (ALA)) and 103 km to the west on the Malaspina Forelands from a station at Yakutat (59°29.86'’ N 139°45.60’ W, Stair & Pennell s.n., 1945 (PH), Hultén 1941-1950, Stair and Pennell 1947). Map 64. Carex williamsii Britt., Williams’ Sedge — NUTZOTIN MOUNTAINS: wet sedge meadow tundra at margin of kettle lake, northwest of Beaver-Horsfeld Creek con- fluence, 1097 m, 62°1' N 141°11’ W, Parker & Gracz 6844, 10 August 1996. This amphiberingean arctic-alpine sedge is rare in the Yukon Territory (Douglas et al. 1981). The collection cited above extends its range 205 km southeast into the Nutzotin Mountains from a station in the Alaska Range (Hultén 1968) and connects the range 155 km to the east in the Yukon Territory (Cody 1996). Map 65. Eriophorum callitrix Cham., Arctic Cotton-grass — CHUGACH MOUNTAINS: moist soil near edge of pond, Martin Creek, 1097 m, 60°56’ N 142°23’ W, Batten & Barker 96-246 (ALA), 96-264, 28 July 1996 (ALA). WRANGELL MOUNTAINS: Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5'’ W, M. Potkin 95-100, 30 July 1995; alpine basin, NW flank of Mt. Drum, 1615 m, 62°4.5’ N 144°45.91' W, C. Meyers 84-23, 4 July 1984; scattered in wet graminoid meadow, Chetaslina Ridge, 1615 m, 61°56.51' N 144°25.93' W, C. Roland 94-360, 17 August 1994. This circumpolar arctic-alpine grass was collected in the Wrangell Mountains at Chitistone Pass (Scott 1968). The localities cited above extend its range 151 km west into the western Wrangell Mountains and 87 km south into the Chugach Mountains. Map 66. Eriophorum viridi-carinatum (Engelm.) Fern., Tassel Cotton-grass — WRANGELL MOUNTAINS: occasional in Sphagnum bog, Lakina River, Chitina River Basin, 792 m, 61°29.49' N 143°25.6’ W, C. Roland 96-672, 24 July 1996. This North American boreal-montane cotton-grass is rare in Alaska (G5 $2) and in the Yukon Territory (Douglas et al. 1981). The specimen cited above connects its distri- bution 259 km to the east in the Anchorage Quad (Muldoon bog, 61°12.22’ N 149°42.97’ W, Duffy & Tande 923, 26 July 1994 (ALA)) with a station 458 km to the southeast. near Haines (Hultén 1968). Map 67. Kobresia sibirica Turcz., Siberian Bog Sedge — NUTZOTIN MOUNTAINS: patchy on upper slope of S- facing bluff, White River, 1219 m, 61°44.3’ N 141°9.5' W, M. Cook 95085, 21 June 1995. THE CANADIAN FIELD-NATURALIST Vol. 116 This species is circumpolar with an arctic-alpine distri- bution. The specimen cited above extends its range 270 km to the southeast into the Nutzotin Mountains from a collec- tion in the Alaska Range (Bear Creek, 63°37.0' N 145°50.0’ W, Batten & Dawe 78-42, 29 June 1978 (ALA)) and connects the range 62 km to the east in the Yukon Territory (Cody 1996). Map 68. Kobresia simpliciuscula (Wahlenb.) Mack., Simple Bog Sedge — MENTASTA MOUNTAINS: co-dominant in hummocky meadow on floodplain, Lost Creek, 1006 m, 62°32’ N 143°9.6’ W, Roland & D’Auria 97-062, 97-073, 26 June 1997. NUTZOTIN Moun- TAINS: mesic graminoid herbaceous meadow between Gold Run and Glacier Creeks, 1493 m, 62°7.42' N 141°52.73' W, M. Cook 8983, 23 July 1989; wet sedge meadow tundra at margin of kettle lake, Beaver-Horsfeld Creek confluence, 1097 m, 62°1’ N 141°11’ W, Parker & Gracz 6843, 6857, 6900, 10 August 1996; sedge meadow, N of Francis Creek, 1234 m, 61°53.78' N 141°8.81’ W, M. Duffy 96-273, 8 August 1996. The collections cited above of this circumpolar arctic- alpine sedge extend its range 136 km south into the Mentasta Mountains and 244 southeast into the Nutzotin Mountains from the Alaska Range (Hultén 1968) and connect its distribution 130 km to the east in the Yukon Territory (Cody 1996). Map 69. Trichophorum pumilum (M. Vahl.) Schinz. & Thell. var. rollandii (Scirpus rollandii Fern.), Rolland’s Leafless-Bulrush — ST. ELIAS MOUNTAINS: wet allu- vium at terminus of Barnard Glacier, St. Elias Mountains, 488 m, 61°5.82’ N 141°57.92' W, Duffy & Barnes 96-036, 5 August 1996. This boreal montane sedge is new to the state of Alaska. The collection cited above represents the western limit of its range and is 191 km west of collections in the Yukon Territory (Cody 1996). It is rare in Alaska (G3T3T4? S1) and in the Yukon Territory (Douglas et al. 1981). Map 70. JUNCACEAE Juncus filiformis L., Thread Rush — CHUGACH MOunrtTAINS: abundant in mud, Granite River, 884 m, 60°44.63' N 142°6.05’ W, C. Roland 96-790, 29 July 1996; scattered at margin of subarctic lowland sedge wet meadow, upper Tebay Lake, 579 m, 61°11’ N 144°24' W, Parker & Gracz 6738, 7 August 1996. ST. ELIAS MOUNTAINS: wet sedge herb meadow tun- dra, Short River, 503 m, 6195.35’ N 141°56.3’ W, Parker & Duffy 6681, 6 August 1996. WRANGELL MOUNTAINS: scattered on sandy beaches and alluvial flats, Fox Farm Lake, 727 m, 62°19.98’ N 144°50.02'’ W, C. Roland 95-260, 25 July 1995; occasional in bare clay soil in graminoid meadow, SE end Boulder Lake, 1036 m, 62°31-277> 144°11.26’ W, M. Cook 95365, 2 August 1995; com- mon along lake shore in open silt, Copper River, 911 m, 62°24.53’ N 143°42.68' W, C. Roland 96-917, 11 August 1996. This rush is circumpolar with a boreal-montane distribu- tion. The collections cited above extend its range southeast 2002. COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 68. Kobresia sibirica 231 tee eee ee ee ———e 232 103 km into the Wrangell Mountains and 330 km into the Chugach Mountains from a station in the Alaska Range (Mile 3.3 Denali Highway, 63°40.0' N 145°33.0' W, L.A. Viereck 8359, 26 July 1967 (ALA)). Map 71. Juncus mertensianus Bong., Merten’s Rush — CHUGACH MOUNTAINS: rocks and saturated silt, lower Bremner River, 488 m, 61°0.81' N 144°14.39’ W, C. Meyers 84-104, 20 July 1984; sand in open willow scrub, Granite River, 884 m, 60°44.32’ N 142°4.06’ W, M. Cook 96616, 7 July 1996; scattered forbs, Short River, 503 m, 61°5.35’ N 141°56.3’ W, Parker & Duffy 6691, 6 August 1996; scattered in moss, 12-Mile.Basin,, 1326 m,.60°50.21' N 142°30.85' W, M. Cook 96599, 7 July 1996; along stream in forb herbaceous vegetation, Falls Creek, 655m: GI°T4.17’ N 144°28.24 WW. TOA. ee: G. Viereck 11044, 7 July 1996; tall willow scrub, Granitic Creek, 1188 m, 61°4.37' N 142°56.79’ W, M. Hoffman s.n., 11 August 1986; in floating peat on lakeshore, Martin Creek, 1097 m, 60°56’ N 142°23’ W, Batten & Barker 96-257, 28 July 1996; abundant in meadow, Canyon Creek, 1035 m, 61°20.17' N 144°19.38'’ W, C. Roland 96-549, 15 July 1996; common in heath, 12-mile Creek, 988 m, 60°50.51’ N 142°23.02’ W, C. Roland 96-765, 7 July 1996; seepage area at margin of subarctic lowland sedge wet meadow, Granite Creek, 701 m, 60°43.81' N 142°30.08' W, Parker & Gracz 6773, 8 August 1996. St. ELIAS MOUNTAINS: herbaceous meadow, Karr Hills, 60°8.66’ N 141°16.44’ W, K. Beck s.n., 12 August 1987. WRANGELL MOUNTAINS: patchy in wet mud, Grant Creek, -853'm, 61°18,16 “N 143°51.87' W, Cook & Losso 96334, 7 July 1996; moist fine sand along rivulet, Skolai Creek, 1341 m, 61°41.5’ N 142°23' W, Batten & Barker 96-129, 25 July 1996; patchy in graminoid-forb meadow, Cheshnina Plateau, 1399 m, 61°48.04’ N 144°6.27’ W, C. Roland 96-869, 7 August 1996. This rush is North American with a cordilleran distribu- tion. The specimens cited above extend its range east 225 km into the Chugach Mountain, 285 km into the south- ern St. Elias Mountains and northeast 211 km into the Wrangell Mountains from a collection in the Valdez Quad (61°12.0’ N 145°47.0' W, L.A. Viereck 8472, 1 August 1967 (ALA)). These localities also connect the range 151 km to the east in the Yukon Territory (Cody 1996). Map 72. LILIACEAE Maianthemum stellatum (L.) Link., Star-flowered Solomon’s Seal — NUTZOTIN MOUNTAINS: Populus balsamifera forest on steep slope, between Sheep and Notch Creeks, 488 m, 62°15.09’ N 141°59.17’ W, D. Morrison 84-41, 19 June 1984. This lily is North American with a boreal-montane dis- . tribution and is rare in Alaska (G5 S2). The specimen cited above extends its range 328 km to the east into the Nutzotin Mountains from a collection in the Anchorage Quad (Long Lake, 61°48.55’ N 148°14.56’ W, M. Cook 3114, 2 June 1998 (ALA)) and 292 km south from collections in the Eagle Quad (Mission Creek, 64°48.0’ N 141°10.0’ W, THE CANADIAN FIELD-NATURALIST Vol. 116 Khokhryakov et al. 6294, 10 July 1981 (ALA)). This locali- ty also connects the distribution 170 km to the east in the Yukon Territory (Cody 1996). Map 73. SALICACEAE Salix commutata Bebb, Undergreen Willow — CHUGACH MOUNTAINS: flooded gravel bar, Martin Creek, 1097 m, 60°56’ N 142°23' W, Batten & Barker 96-262, 28 July 1996; scattered in wet silt with Equisetum, Copper/Bremner River confluence, 152 m, 60°56.95’ N 144°41.79’ W, C. Roland 96-605, 17 July 1996. GULF OF ALASKA BASIN: graminoid herbaceous meadow, Alder Stream, 59°44.22' N 140°22.95'’ W, M. Cook 87-94, 16 August 1987; horsetail bog, Cape Sitkagi, 61 m, 59°47.52’ N 140°56.51' W, M. Cook 87-99, 20 August 1987; wet sedge marsh, Esker Stream, 59°54.19' N 139°46.27’ W, K. Beck s.n., August 1987. The specimens cited above of this North American cordilleran willow extend its range 148 km to the east into the Chugach Mountains from a station on the Copper River near the Bremner River (Hultén 1968) and connect the range 90 km to the southeast near Yakutat (Harlequin Lake, 59°24.0' N 139°00.0’ W, L.A. Viereck s.n., 7 July 1965 (ALA)). Map 74. Salix rotundifolia Trautv. ssp. dodgeana Rydb. — MENTASTA MOUNTAINS: limestone scree, Soda Lake, 1173 m, 62°32.36’ N 142°53.98’ W, C. Roland 94- 251, 21 July 1994; common in rock crevices and stony tundra, Trail Creek, 1615 m, 62°36.05’ N 143°17.63’ W, C. Roland 95-018, 7 June 1995; com- mon in calcareous gravel, Totschunda Creek, 1280 m, 62°27.63' N 142°12.44’ W, C. Roland 96-290, 24 June 1996. NUTZOTIN MOUNTAINS: limestone gravels, Baultoff Creek, 1707 m, 62°9.13’ N 141°14.51’ W, C. Roland 94-169, M. Cook 94187, 28 June 1994. WRANGELL MOuNTAINS: blocky stone slope, Devil’s Mountain, 1829 m, 62°27.97' N 142°54.97’ W, M. Duffy 91098, 30 July 1991; alpine tundra, Nabesna Glacier moraine, 1798 m, 61°56.6’ N 143° W, M. Duffy 92116, 26 June 1992; scree slope, ridge S of Solo Flats, 1935 m, 61°28.41’ N 141°30.7' W, M. Duffy 92177, 8 July 1992; talus slope, Lime Creek ridge, 1646 m, 61°46.25’ N 141°50.36’ W, M. Cook 94110A, 21 June 1994; rubble slope, NE slopes of Devil’s Mountain, 1530 m, 62°25.62'’ N 142°53.93’ W, C. Roland 96-357, 3 July 1996. This willow is North American with a cordilleran distri- bution. The specimens cited above extend its range 193 km southeast into the Wrangell Mountains from a station in the Alaska Range (Argus 2000) with intermediary stations in the Mentasta and Nutzotin Mountains. Map 75. Salix setchelliana Ball, Setchell’s Willow — CHUGACH MOUNTAINS: rare in wet silt on floodplain, Bremner River, 152 m, 60°56.95' N 144°41.79' W, C. Roland 96-611, 17 July 1996; dunes at mouth of Bremner River, 69 m, 60°59.67’ N 144°34.63’ W, L. A. & E. G. Viereck 11050, 7 July 1996. MENTASTA MOUNTAINS: common in sandy areas of gravel bar, Totschunda Creek, 725 m, 62°26.94’ N 42°40.8’ W, 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 71. Juncus filiformis 72. Juncus mertensianus 233 ee me a 234 C. Roland 96-252, M. Cook 96250, 23 June 1996. WRANGELL MOUNTAINS: scattered on sandy river bar, Bond Creek, 783 m, 62°19.42' N 142°51.82' W, C. Roland 96-176, 19 June 1996; scattered on stabilized river terrace, confluence of Jacksina and Nabesna Rivers 759 mm, 62°21 .51) IN 425287” Wi iG. Roland 96-198, M. Cook 96222, 21 June 1996. This Alaksa- Yukon endemic willow was collected at the head of the Chitina River (H.M. Laing 224, 1925, 224 (CAN) (Porsild 1939; Hultén 1941-1950)) and is rare in Alaska (G3G4 S3) and in the Yukon Territory (Douglas et al. 1981). The new localities cited above extend its range 183 km west into the Chugach Mountains and 184 km northwest into the Mentasta Mountains. These stations con- nect its range 198 km to the west in the Anchorage Quad (Matanuska, 61°32.0' N 149°14.0' W, J.P. Anderson 884, 10 July 1931 (ALA)) and 139 km to the north on the Delta River (63°33.0’ N 145°50.0’ W, G.W. Argus 13398, 15 August 1989 (ALA)) with its range 92 km to the east in the Yukon Territory (Cody 1996). Map 76. Salix stolonifera Cov., Stoloniferous Willow — ST. ELIAS MOUNTAINS: moist gravels, Short River, 503 m, 61°5.35’ N 141°56.3' W, Parker & Duffy 6684, 6 August 1996. This willow is endemic to Alaska and British Columbia and has a Pacific Coastal distribution. The specimen cited above extends its range 200 km northeast into the Chitina River basin from collections near Cordova (Argus 2000) and connects the range 229 km to the southeast at Cordova (Argus 2000). Map 77. POLYGONACEAE Rumex acetosa L. ssp. alpestris (Scop.) Love, Garden Sorrel — NUTZOTIN MOUNTAINS: dry, open shale surrounded by birch/willow shrub tundra, vic. Wiki Peak, 1433 m, 61°53.9’ N 141°9.51’ W, C. Roland 94-114, 22 June 1994; several patches in bare mineral soil surrounded by Dryas moss tundra, Horsfeld Creek, 1768 m, 62°2.88'’ N 141°13.18' W, M. Cook 94162, 24 June 1994; several large popula- tions on SW-facing scree slope, Horsfeld Creek, 1768 m, 62°2.88' N 141°13.18’ W, C. Roland 94- 129A, 24 June 1994; few in moist fine mud of exposed lake shore, Braye Lakes, 1097 m, 62°3’ N 141°6’ W, Parker & Gracz 6865, 11 August 1996; few in sagebrush-grass vegetation on bluffs of old river terraces above Beaver Creek, 1097 m, 62°1’ N 141°11' W, Parker & Gracz 6897, 12 August 1996. This cordilleran species is rare in the Yukon Territory (Douglas et al. 1981). The collections cited above extend its range 607 km east from Lake Clark (Trail Creek, 60°50.0' N 153°49.0' W, P. Caswell 96257, 21 July 1996 (ALA)) and connect the distribution 151 km to the east in the Yukon Territory (Cody 1996). Map 78. Rumex beringensis Yurtsev & Petrovsky (R. graminifolius DC), Bering Sea Dock — St. ELIAS » MOUNTAINS: scattered in volcanic ash deposit, Cub Creek, 1280 m, 61°37.27'’ N 141°9.75' W, M. Cook 94182A, 28 June 1994; occasional in bare soil on knoll in dry tundra, Mt. Natazhat, 1716 m, 61°35.38' N 141°1.83’ W, C. Roland 95-065, 19 June 1995. THE CANADIAN FIELD-NATURALIST Vol. 116 WRANGELL MOUNTAINS: bare area under boulder, West Glacier, 1524 m, 62°18.31’ N 143°52.37’' W, M. Duffy 92021, 23 June 1991, M. Duffy 92267, 9 September 1992; patchy on S-facing scree slope, NE slope of Mt. Drum, 1433 m, 62°8.83’ N 144°30.18’ W, M. Cook 94058, C. Roland 94-075, 11 June 1994; rare on alluvium below cinder cone, north slope of Mt. Sanford, 1722 m, 62°23.4’ N 144°15.9’ W, C. Roland 94-009, 6 June 1994; scattered in bare sandy mineral soil, Chetaslina plateau, 1615 m, 61°56.51’ N 144°25.93’ W, C. Roland 94-343, 15 August 1994; M. Cook 94479, 16 August 1994; occasional in moist bare soil of solifluction lobes, Black Mountain, 1481 m, 62°20.85’ N 143°44.9' W, Cook & Beck 95140, 7 July 1995; abundant in slumping lobes of organic-rich soil around rodent dens, volcanic ridge 6.4 km east of Snyder Peak, 1524 m, 62°4.47' N 144°30.51' W, C. Roland 96- 379, 5 July 1996; scattered throughout alpine basin, upper Dadina River, 1344 m, 62°3.95’ N 144°25.18' W, M. Cook 96267, 7 July 1996; scattered in moist gravel of alpine lake shore, between Klawasi and Nadina Rivers, 1042 m, 62°1.95’ N 144°52.49’ W, M. Cook 96273, 7 July 1996; patchy in bare sandy areas along edge of cliff, Cheshnina Plateau, 1399 m, 61°48.04' N 144°6.27' W, C. Roland 96- 850, 6 August 1996; M. Cook 96700, 8 August 1996. This amphiberingain species with a Pacific coastal distribution, is rare in Alaska (G3 S3). It had been collected by David F. Murray at Sheep and Guerin Glaciers in the St. Elias Mountains (Murray 1971). The specimens cited above extend its range 201 km into the western Wrangell Mountains. These collections document the eastern limit of its range and are disjunct from the closest locality 507 km to the southwest at Lake Clark (Tanalian Point, 60°12.0’ N 154°19.0' W, D.R. Hunt 44, 11 June 1993 (ALA)). Map 79. PORTULACACEAE Claytonia tuberosa Pall., Tuberous Springbeauty — NUTZOTIN MOUNTAINS: open area within closed low birch scrub, vic. Rock Lake, 1113 m, 61°49.6’ N 141°20’' W, M. Duffy 91090, 22 July 1991; rare in subalpine meadow within open spruce forest, Carden Hills, 1311 m, 62°18.44’ N 141°11.55’ W, C. Roland 94-134A, 25 June 1994; scattered in bare, wet ash deposit, Flat Lake, 1323 m, 61°58.97’ N 141°41.25’ W, C. Roland 95-046, 17 June 1995; scattered in wetlands along Carl Creek, 1402 m, 62°2.81' N 141°34.65’ W, M. Cook 3153, 2 July 1998. This species is amphiberingian with a boreal-montane distribution. The collections cited above extend its range 158 km south into the Nutzotin Mountains from a collec- tion in the Tanacross Quad (Mt. Fairplay, 63°40.0’ N 142°13.0' W, G. Smith 2426, 30 June 1954 (ALA)) and connect the range 140 km east in the St. Elias Mountains of the Yukon Territory (Cody 1996). Map 80. Montia bostockii (A.E. Porsild) Welsh (Claytonia bostockii Pors.), Bostock’s Minerslettuce — CHUGACH MOUNTAINS: sedge-Dryas tundra, Virgin Gulch, 1539 m, 61°15.52’ N 142°31.09’ W, M. Protti 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 76. Salix setchelliana 255 236 9104, 17 July 1991. MENTASTA MOUNTAINS: numer- ous in moist sedge/birch meadow, Big Grayling Lake, 1036 m, 62°32.93' N 143°143.08’ W, G. Dodge s.n. June 1993; scattered in mesic Dryas tundra, ridge E of Soda Lake, 1173 m, 62°32.36' N 142°53.98’ W, M. Cook 94307, 21 July 1994; occasional in open Picea mariana muskeg, Lost Creek floodplain, 1006 m, 62°32’ N 143°9.6' W, Roland & D’Auria 97-074, 26 June 1997. NUTZOTIN MOUNTAINS: open shrub thicket, confluence of Notch Creek and Chisana River, 1097 m, 62°10.5’ N 142°5.5’ W, J. Bolivar 84-48, 25 June 1984; moist sedge meadow, Solo Flats, 1402 m, 61°50.67' N 141°47.05’ W, M. Duffy 92153, 7 July 1992; rare in low areas between dunes, Beaver Lake, 1341 m, 62°2.61’ N 141°48.39’ W, M. Cook 94198, 29 June 1994; scattered in moss along stream bank, volcanic plateau between Bryan and Willow Creeks, 1829 m, 61°58.97' N 141°50.03’ W, M. Cook 95055B, 18 June 1995; few in saturated moss, Braye Lakes, 1097 m, 62°2' N 141°9’ W, Parker & Gracz 6870, 11 August 1996; saturated moss, open birch-willow serub, Ptarmigan Lake, 1097)m,.61750:03" N 141°13.75’ W, Duffy & Barnes 96-211, 8 August 1996. St. ELIAS MOUNTAINS: open tundra muskeg, McColl Ridge, 1341 m, 61°12.36’ N 142°39.64' W, J. Bolivar 84-106, 15 July 1984. WRANGELL MOUNTAINS: tussock muskeg ridgetop, between Grizzly Lake and Jacksina Creek, 1890 m, 62°14.51' N 143°17.46' W, J. Bolivar 84-115, 23 July 1984; Dryas-sedge tundra, NW slope of Mt. Sanford, 1296 m, 62°21.17’ N 144°24.52' W, M. Cook 93125, 9 June 1993; Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5’ W, M. Potkin 95-067, 28 July 1995; occasional in wet swale, bench between Monte Cristo Creek and Nabesna River, 1021 m, 62°14.07’ N 142°56.85’ W, C. Roland 96-139, 18 June 1996. This arctic-alpine Alaska-Yukon endemic was collected by D. F. Murray and R. W. Scott at Chitistone Pass in the Wrangell Mountains (Murray 1968, Scott 1968). The speci- mens cited above extend its range 152 km west into the Wrangell Mountains, 61 km south into the St. Elias Mountains, 114 km northwest into the Mentasta Mountains and 62 km north into the Nutzotin Mountains. The closest localities are 77 km to the north in the Nabesna Quad (Tetlin Mountains, 62°41.0' N 142°46.0' W, K. Teare 1610, 14 July 1983 (ALA)) and 58 km to the east in the Yukon Territory (Cody 1996). It is rare in Alaska (G3 S3) and in the Yukon Territory (Douglas et al. 1981). Map 81. CARYOPHYLLACEAE Arenaria capillaris Poir., Beautiful Sandwort — CHUGACH MOUNTAINS: talus slope and Dryas tundra, Hundell Creek, 1615 m, 61°36.75’ N 144°42.2’ W, M. Cook 93465, 28 August 1993; dry S-facing slope, Towhead Mountain, 1433 m, 61°3.21’ N 142°39.8’ W, Duffy & Barnes 96-176, 8 August 1996; Dryas- lichen tundra, Five-Mile Creek, 1219 m, 61° 32.29'N 144 33.11'W, M. Cook 3539, 15 July 2000. This amphiberingian cordilleran species, rare in the Yukon Territory, was collected in the vicinity of the Park by J.P. Anderson along the Slana-Tok Highway in 1935 THE CANADIAN FIELD-NATURALIST Vol. 116 (Hultén 1940-1951), Charles W.H. Heidemann in 1908 near Copper Center (Hultén 1940-1951) and at Tazlina Glacier (61°45.0’ N 146°30.0’ W, L. A. & E.G. Viereck 2191, 19 July 1957 (ALA)). The specimens cited above extend its range 111 km east in the Chugach Mountains. Map 82. Arenaria longipedunculata Hult., Longstem Sand- wort — ST. ELIAS MOUNTAINS: wet alluvium of grav- el bar, Barnard Glacier terminus, Chitina River, 488 m, 61°5.82’ N 141°57.92' W, Duffy & Barnes 96-028, 8 August 1996. This North American cordilleran species is rare in Alaska (G3G4Q S3) and of limited distribution in the Yukon Territory (Cody 1996). The specimen cited above extends its range 536 km southeast into the St. Elias Mountains from a locality near Healy (A.E. Porsild 339, 1926 (CAN), Hultén 1941-1950) and connects the distribu- tion 212 km to the southwest in the Yukon Territory (Cody 1996). Map 83. Cerastium regelii Ostenf., Regel’s Chickweed — WRANGELL MOUNTAINS: mesic sedge grass meadow tundra, Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5’ W, Potkin & Leggett 95-085, 29 July 1995 (ALA). This species is circumpolar with an arctic-alpine distribu- tion and is rare in Alaska (G4 S2S3). Our distribution map reflects the combination of C. jenisejense with C. regelii by Heide et. al. (1990). The specimen cited above is 1071 km to the east of a collection made by Eric Hultén at Cape Newenham in western Alaska and 417 km south of a station in the Steese-White Mountains (Hultén 1968). Map 84. Minuartia biflora (L.) Schinzl. & Thell., Mountain Stitchwort — CHUGACH MOUNTAINS: moist bouldery solifluction lobe, East Fork Kiagna River, 1487 m, 60°59.5' N 142°1’ W, Batten & Barker 96-311, 29 July 1996 (ALA); scattered in Rhacomitrium of dry tundra, Nerelna Creek plateau, 1372 m, 61°26.66’ N 144°17.69' W, M. Cook 96309, Roland & D’Auria 96-411, 8 July 1996. MENTASTA MOUNTAINS: scat- tered in mesic dwarf willow-Dryas-moss communi- ty, Lost Creek, 1722 m, 62°36.45’ N 143°12.03’ W, M. Cook 95169, 7 July 1995. NUTZOTIN MOUNTAINS: occasional in bare organic soil between boulders at base of scree slope, 1585 m, 62°25.78’ N 142°22.11' W, M. Cook 95107, 28 June 1995; moist sandy soil with seral herbs on well-vegetated SE-facing alpine colluvium, Horsfeld Creek, 1128 m, 62°2’ N 141°11’ W, Parker & Gracz 6914, 13 August 1996. ST. ELIAS MOUNTAINS: occasional in lush, steep, S-fac- ing meadow, Mt. Natazhat, 1716 m, 61°35.38’ N 141°1.83’ W, C. Roland 95-068, 20 June 1995; few in bare organic soil on rock moraine, Mt. Chitina, 2073 m, 60°57.74' N 141°17.33' W, M. Cook 95224, C. Roland 95-195, 14 July 1995. WRANGELL MOUNTAINS: moist gravel, sparsely vegetated, SE slope of Fredrika Mountain, 1859 m, 61°43.8’ N 142°6.46' W, M. Duffy 91092, 2 July 1991; rare in mesic stony area, Cone Ridge, 2073 m, 62°8.21’ N 143°18.47' W, C. Roland 94-299, 25 July 1994; rare in mesic, mossy area of N-facing gravel slope, 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 80. Claytonia tuberosa 237 - CC OO OC --:C + --—-—--— or OS — 3 - 238 THE CANADIAN FIELD-NATURALIST Vol. 116 84. Cerastium regelii 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 88. Sagina saginoides 239 i ee _—_——_—- lc Ol PS eS = 240 Crystalline Hills, 1585 m, 61°23.59' N 143°31.85' W, C. Roland 95-249, 18 July 1995; dry tundra on ridge Nikolai Pass, 1280 m, 61°26’ N 142°40’ W, Batten & Barker 96-002, 23 July 1996 (ALA); rare in mossy, stony saturated soil of seep, Hasen Creek, 1835 m, 61°34.06’ N 142°18.33’ W, C. Roland 96- 643, 23 July 1996; scattered in sedimentary gravel scree on lower third of slope, West Fork Mill Creek, 1241 m, 619°33.33’ N 143°28.35' W, M. Cook 96519, 24 July 1996; occasional in moss between gravel at base of slope, Lakina Glacier, 1219 m, 61°33.64' N 143°19.01' W, M. Cook 96545, 24 July 1996; moist muddy scree, Nikolai Mine, 1695 m, 61°27’ N 142°39' W, Batten & Barker 96-O089B, 24 July 1996 (ALA); moist gravelly clay, VABM Sentinel west of Nizina Glacier, 1829 m, 61°39’ N 142°32’ W, Batten & Barker 96-110A, 24 July 1996 (ALA); scree, ridge W of Nikolai Pass, 1372 m, 61°26.5'’ N 142°43’ W, Batten & Barker 96-161A, 26 July 1996. This circumpolar arctic-alpine species, which is rare in Alaska (GS $2), was previously known from three locali- ties in the Wrangell Mountains: Chitistone Pass (61°37' N 142°3' W, R. Scott 1630, 1867, 2214 (MICH), Scott 1968), Skolai valley (61°37’ N 141°58’ W, D. F. Murray 719 (CAN), Murray 1968) and Bonanza Ridge (61°30’ N 142°51' W, Nordell & Schmitt 95, 416b, 450, (LD & ALA), Nordell and Schmitt 1968). The new collections cited above extend the range of this species 124 km north into the Mentasta Mountains, 64 km north into the Nutzotin Mountains, 123 km southwest into the Chugach Mountains, 75 km northwest in the Wrangell Mountains and 105 km southeast into the St. Elias Mountains. These collections join the range 152 km to the northeast in the Mt. Hayes Quad (Tangle Lakes, 63°03.0’ N 146°01.0’ W, G. Smith 2033, 20 August 1953 (ALA)) with the range 204 km to the east in the Yukon Territory (Cody 1996). Map 85. Minuartia dawsonensis (Britt.) Mattf., Rock Stitchwort — CHUGACH MOUNTAINS: Copper River bluff below Gulkana airport, 457 m, 62°10.45' N 145°24.3' W, D. Taylor 8536, 11 July 1984. Nut- ZOTIN MOUNTAINS: Black Spruce woodland, Tanana River lowlands, 2900 m, 62°25.09’ N 141°34.33’ W, M. Duffy 91010, 19 June 1991; scattered on shore, Braye Lakes, 1097 m, 62°2’ N 141°9’ W, Parker & Gracz 6883, 11 August 1996. St. ELIAS MOUNTAINS: shore of Mt. Chitina lake, 823 m, 60°57.74’ N 141°17.33' W, M. Cook 95260, 17 July 1995. WRANGELL MOUNTAINS: spruce-aspen woodland, small patch on trail, 3 mi E of Strelna Creek, 619 m, 61°33.98' N 144°1.56’ W, M. Cook 3137, 17 June 1998 (ALA); mixed woodland floodplain gravel bar, Kennicott River, 427 m, 61°26.38'’ N 142°57.64’ W, M. Cook 3515, 30 September 1998 (ALA). This species is North American with a boreal-montane distribution. The specimens cited above extend its range 160 km into the southern Wrangells, 160 km southeast into the Nutzotin Mountains and 237 km southeast into the St. Elias Mountains from a station near Mentasta Pass (Hultén 1968). These collections connect the distribution in the Alaska Range with the distribution 129 km to the east in the Yukon Territory (Cody 1996). Map 86. THE CANADIAN FIELD-NATURALIST Vol. 116 Minuartia stricta (Sw.) Hiern, Rock Sandwort — NUTZOTIN MOUNTAINS: wet sedge meadow tundra at margin of kettle lake, vic. Horsfeld Creek, 1097 m, 62°1' N 141°11’ W, Parker &(Gracz 6Ga42) 10 August 1996. This circumpolar arctic-alpine species was known from Bonanza Ridge in the Wrangell Mountains (61°30’ N 142°51' W, Nordell & Schmitt 256, 463 (LD & ALA) (Nordell and Schmitt 1977)) and a station near Slana (Hultén 1968). The specimen cited above extends its range 104 km east into the Nutzotin Mountains. Map 87. Sagina saginoides (L.) Karst., Arctic Pearlwort — CHUGACH MOUNTAINS: occasional in wet mud, Grant Creek, Cook & Losso 96339, 8 July 1996; shady steep bouldery soil, Goat Creek, 1487 m, 60°59.5’ N 142°1' W, Batten & Barker 96-312B, 29 July 1996 (ALA); lush meadow, 12-Mile Basin, 1326 m, 60°50.21' N 142°30.85’ W, C. Roland 96-783, 29 July 1996. COPPER RIVER BASIN: sandy riverbed, Gulkana Airport, 381 m, 62°10.5' N 145°24.57' W, K. Teare 1658A, 8 June 1984. GULF OF ALASKA BASIN: open willow scrub, 60°5.43’ N 141°25.51’ W, M. Duffy 92255, 22 August 1992. WRANGELL MOUNTAINS: small patches of fine soil surrounded by cobbles, Skolai Creek, 1341 m, 61°41.5’ N 142°23’ W, Batten & Barker 96-125, 25 July 1996 (ALA). This species is circumpolar with a widespread distribu- tion. The localities cited above extend its range 150 km east into the Wrangell Mountains, 168 km southeast into the Chugach Mountains and 268 km southeast into the southern St. Elias Mountains from stations near Copper Center and Willow Creek along the Richardson Highway (Hultén 1968). Map 88. Silene involucrata (Cham. & Schlecht.) Bocquet ssp. involucrata (Melandrium affine J. Vahl), Arctic Catchfly — CHUGACH MOUNTAINS: rock outcrop, Towhead Mountain, 1433 m, 61°3.21’ N 142°39.8’ W, Duffy & Barnes 96-154, 8 August 1996. MENTASTA MOUNTAINS: rare in understory of dry Populus balsamifera forest on steep, S-facing slope, Lost Creek , 1097 m, 62°33.3’ N 143°9' W, Roland & D’Auria 97-035, 26 June 1997. NUTZOTIN MounrtTaAINs: S-facing scree and N-facing graminoid- forb tundra, vic. Ptarmigan Creek, 1494 m, 61°32.89’ N 141°3.28' W, M. Cook 92480, 9 July 1992; rare in meadow tundra, Carden Hills, 1311 m, 62°18.44’ N 141°11.55' W, C. Roland 94-134B, 25 June 1994; occasional on S-facing talus slope, Rocker Creek, 1433 m, 61°54.8’ N 141°2.03’ W, Cook & Roland 94139, 23 June 1994. WRANGELL MOuNnNTAINS: N-facing scree slope, ridge between Lime Creek and Solo Flats, 1707 m, 61°28.41’ N 141°29.44’ W, M. Cook 92500, 92109, 7 July 1992; Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24’ W, Moran & Roland 95-39, 1 July 1995; few on S-fac- ing bluff, Ellis Lake, 1219 m, 62°16.6’ N 142°56’ W, M. Cook 3032, 21 June 1997. This species is circumpolar with an arctic-alpine distri- bution. It had been collected in the Wrangell Mountains 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 91. Silene repens 92. Silene uralensis ssp. uralensis 241 - --+---ooOoCroOo Orr oe SO 242 (Bonanza Ridge, 1219 m, 61°30’ N 142°51’ W, Nordell and Schmitt 622, 1967 (ALA, LD) (Nordell and Schmitt 1977; Hultén 1968)) and in the St. Elias Mountains (head of Chitina River, H.M. Laing 228, 1925 (CAN) (Porsild 1938; Hultén 1941-1950)). The specimens cited above extend its range 57 km to the east into the Chugach Mountains, 84 km to the northwest in the Wrangell Mountains, 122 km to the northwest into the Mentasta Mountains, 126 km north into the Nutzotin Mountains and 68 km north into the St. Elias Mountains. These collections connect its range 186 km to the northwest in the Alaska Range (Mt. Hayes Quad: 63°37.0’ N 146°43.0’ W, L.J. Palmer 665, 15 June 1941 (ALA)) with the range 116 km to the east in the Yukon Territory (Cody 1996). Map 89. Silene menziesii Hook., Menzies’ Campion — ST. ELIAS MOUNTAINS: rare in bare soil around rodent burrows in alpine forb herbaceous meadow, ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94’ N 141°6.75' W, C. Roland 95-180, 12 July 1995; C. Roland 95-184, 13 July 1995; M. Cook O527 7,43 July 19935; This North American cordilleran species was collected on the upper Chitina River by Hamilton M. Laing in 1925 (#61, 62 (CAN)). Porsild identified this material as S. williamsii Britt. (Porsild 1938), but Hitchcock and Maguire (1947) re-determined it as S. menzeisii. This determination also fits the distribution pattern better than S. williamsii, which is an Alaska-Yukon endemic, species which are more common in the mountains of the Alaska Range in the Park. The station in Hultén (1968) for Laing’s collection on the upper Chitina River is near Hubrick’s Landing where Laing had his base camp while the expedition was climbing Mt. Logan from May 18 to July 6 (Lambart 1926a, 1926b). However, he surveyed from Hubrick’s Landing up river to Logan and Walsh Glaciers, so we cannot be certain where this collection was made. The collections in the Park are 403 km southeast of the closest station in Alaska (Talkeetna Mountains, 62°55.0' N 149°40.0' W, P. Wagner 74, 4 July 1988 (ALA)) and 175 km west of collections in the Yukon Territory (Cody 1996). Map 90. Silene repens Patrin. — MENTASTA MOUNTAINS: occa- sional in dry, rubbly areas on S-facing bluff, Caribou Creek, 975 m, 62°35.1' N 143°28.1’ W, Roland & D’Auria 97-011, 26 June 1997; occasional on S-fac- ing dry forb herbaceous slope, Devil’s Mountains, 942 m, 62°24.95' N 142°54.86' W, M. Cook 96236, C. Roland 96-223, 22 June 1996; scattered in organic soil with lichens on rock outcrops on ridge, Devil’s Mountains, 1530 m, 62°25.62' N 142°53.93' W, M. Cook 96281, 7 July 1996; scattered in greenstone rub- ble, limestone ridge between Soda and Platinum Creeks, 1494 m, 62°31.2’ N 142°54.03' W, C. Roland 95-142, 5 July 1995; scattered along south side of ridge on exposed rock outcrops and gravel areas, between Totschunda and Platinum Creeks, 1494 m, 62°27.82' N 142°46.27' W, M. Cook 3158, 9 July 1998. NUTZOTIN MOUNTAINS: alpine scree slopes, Ptarmigan Lake, 1234 m, 61°53.78’ N 141°8.81' W, J. Barnes 96-261, 12 August 1996; rock slope, Rocker Creek, 1615 m, 62°9.56’ N 145°0.76' W, M. Duffy 92186, 9 July 1992; rare on W-facing open low birch THE CANADIAN FIELD-NATURALIST Vol. 116 scrub slope; Horsfeld Creek, 1128 mpia2-2 ah 141°11’ W, Parker & Gracz 6906, 13 August 1996; occasional in steppe and open poplar woodland, Rock Lake, 1119 m, 61°48.7’ N 141°16.57’ W, C. Roland 96-040, 5 June 1996; common in talus meadow, Wiki Creek, 1411 m, 61°54.46' N 141°6:68% Woe Roland 96-107, 11 June 1996; scattered in Dryas-low scrub, Alder Creek, 1554 m, 62°28.44’ N 142°15.06’ W, M. Cook 95128, 29 June 1995; rare on SE-facing subalpine meadow, Carden Hills, 1311 m, 62°18.44’ N 141°11.55’ W, C. Roland 94-141A, 26 June 1994; S-facing bluff, White River between Cub and Traver Creeks, 1219 m, 61°44.3’ N 141°9.5' W, C. Roland 95-074, 21 June 1995; few scattered in unstable gravel and cobble sized scree, rhyolite knob, Wiki Basin, 1524 m, 61°54.77' N 141°11.05’ W, M. Cook 3184, 15 July 1998; scattered on gravel scree slope, Carl Creek, 1791 m, 62°3.14’ N 141°35.06’ W, M. Cook 3148, 1 July 1998; patchy on boulder slope, Carl Creek, 1590 m, 62°2.69’ N 141°33.97' W, M. Cook 3150, 2 July 1998. WRANGELL MOUNTAINS: few in bare mineral soil, Nabesna River bluff, 1106 m, 62°15.05’ N 142°54.68". W, M. Cook Gane Roland 96-171, 19 June 1996. This speices is amphiberingean with a boreal-montane distribution. It was collected on the Sanford River (W.L. Poto 103, 1902 (US) (Poto 1902*, Hultén 1943)) and the Nabesna River (Schrader & Hartman 61, 1902 (US) (Hultén 1943)). However, these stations do not appear in Hultén’s 1968 flora. The specimens cited above extend its range 71 km into the Mentasta Mountains and 181 km east into the Nutzotin Mountains from the collections on the Sanford and Nabesna Rivers and 202 km south from stations in the Tanana Valley (Hultén 1968). Map 91. Silene uralensis (Rupr.) Bocquet ssp. uralensis (Melandrium macrospermum Pors., S. macrosperma (Porsild) Hultén), Largefruit Catchfly — CHUGACH MOounTAINs: SE-facing ridgetop lichen tundra, vic. Chakina River, 1768 m, 61°6.19’ N 142°55.03’ W, C. Roland 94-184C, 8 July 1994. This species is circumpolar with an arctic-alpine distri- bution. The specimen cited above extends its range 410 km to the southeast into the Chugach Mountains from a station in the Healy Quad (Sable Pass, 63°34.0'’ N 149°40.0’ W, A. Murie s.n., 19 July 1964 (ALA)) and connects the range 148 km to the east in the Yukon Territory (Cody 1996). Map 92. Silene williamsii Britt. (Britt.) Hult. (S$. Hook. ssp. williamsii (Britt.) Hult.), Williams Campion — MENTASTA MOUNTAINS: occasional in seral herb vege- tation on Nabesna River floodplain at Totschunda Creek, 725 m, 62°26.94' N 42°40.8' W, M. Cook 96249, C. Roland 96-253, 23 June 1996. WRANGELL MOoUunrTaAINS: stabilized river terrace with high lichen cover, 2 mi SE of Whitham Lake, 817 m, 62°18.96’ N 142°54.07' W, C. Roland 96-197, 20 June 1996; occasional in sandy areas of river terrace with patchy open low shrubs and mixed forbs, Nabesna River at Jacksina River, 759 m, 62°21.51' N 142°52.87’ W, 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 243 M. Cook 96221, 6 June 1996; scattered in cobbles and sand on river terrace, Copper River, across from Black Mountain, 945 m, 62°19.02’ N 143°47.44’ W, C. Roland 96-904, 10 August 1996. This Alaska-Yukon endemic was collected by F. W. Went in the vicinity of Copper Center just outside the Park in 1934 (Hultén 1943). The specimens cited above extend its range 151 km northeast into the Mentasta Mountains and connect the range 99 km to the north in the Tanacross Quad (Slippery Rock Creek, 63°21.0’ N 143°29.0' W, L.A. Spetzman 721, 19 July 1957 (ALA)) with the station at Copper Center. This species is rare in the Yukon Territory (Douglas et al. 1981). Map 93. Stellaria alaskana Hult., Alaska Starwort — MENTASTA MOUNTAINS: limestone slope, upper Trail Creek, 1341 m, 62°37.09' N 143°16.06'’ W, M. Potkin 95-042, 26 July 1995; scattered at base of NE-facing limestone scree slope, headwaters of Lost Creek, 1722 m, 62°36.45’ N 143°12.03' W, M. Cook 95176, 7 July 1995; limestone slope, upper Trail Creek, 1341 m, 62°37.09’ N 143°16.06’ W, A. Leggett 95-052, 7 July 1995; rare on rock outcrops, Devils’s Mountain, 1530 m, 62°25.62’ N 142°53.93’ W, C. Roland 96-351, 3 July 1996. NUTZOTIN MOUNTAINS: scattered in steep, unstable granitic rub- ble, Carl Creek, 1920 m, 62°3.52’ N 141°36.27' W, C. Roland 95-022, 13 June 1994; moist scree on N- facing slope, Baultoff Creek, 1707 m, 62°9.13’ N 141°14.51’ W, M. Cook 94177A, 27 June 1994; scat- tered underneath rocks in moist loamy sand, Klein Creek, 1747 m, 62°2.29’ N 141°19.88’ W, M. Cook 95012, 15 June 1995; scattered at base of scree slope, alpine basin at headwaters of Stone Creek, 1585 m, 62°25.78' N 142°22.11’ W, M. Cook 95104, 27 June 1995; scattered at base of rock outcrops, headwaters of Alder Creek, 1554 m, 62°28.44’ N 142°15.06’ W, M. Cook 95138, 30 June 1995. Sr. ELIAS MOUNTAINS: scattered on NE-facing unstable volcanic scree slope, Mt. Natazhat, 1716 m, 61°35.38' N 141°1.83’ W, M. Cook 95075, C. Roland 95-071, 20 June 1995. WRANGELL MOunrTAINS: N-facing scree slope, Lime Creek ridge, 1707 m, 61°47.2' N 141°49.06’ W, M. Cook 92498, 7 July 1992; S-facing granite boulder slope, Cooper Pass, 1942 m, 62°17.16’ N 142°31.44’ W, C. Roland 94-285B, 24 July 1994; scattered on upper slope of cinder cone, Cone Ridge, 2073 m, 62°8.21' N 143°18.47' W, M. Cook 94378, C. Roland 94-315, 25 July 1994; rare in tundra, Crystalline Hills, 1585 m, 61°23.59' N 143°31.85' W, C. Roland 95-251, 18 July 1995; scattered on E-facing boulder slope, Fish Creek, 1067 m, 62°16.92' N 142°58.99' W, M. Cook 95318, 27 July 1995; Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5' W, Leggett & Potkin 95-073, 95-084, 28 July 1995; scattered on scree slope, Black Mountain, 1481 m, 62°20.85' N 143°44.9' W, M. Cook 95145, 7 July 1995; steep N-facing scree, Nikolai Pass, 1280 m, 61°26’ N 142°40’ W, Batten & Barker 96-028, 23 July 1996; scree near rocky ridge, VABM Sentinel, west of Nizina Glacier, 1829 m, 61°39’ N 142°32’ W, Batten & Barker 96-124, 25 July 1996 (ALA); scattered in rhyolite scree on SE- facing slope Kuskulana Pass, 1545 m, 61°33.72’ N 143°39.7' W, C. Roland 96-708, 26 July 1996; occa- sional in sandy soil on S-facing slope, 6 km east of Snyder Peak, 1524 m, 62°4.47’ N 144°30.51’ W, C. Roland 96-376, 5 July 1996; occasional in Rhacomitrium at head of scree slope, Kuskulana Pass, 1545 m, 61°33.72’ N 143°39.7’' W, M. Cook 96549, 7 July 1996; rare along gravel creekbed, upper Skookum trail, 1563 m, 62°25.83'N 143°6.91'W, 1572 m, M. Cook 3533, 7 July 1999; scattered on scree slope, Grizzly Lake, 1312 m, 62°14.05'N 143°20.98'W, J. Allen s.n., 27 July 2000. This Alaska-Yukon endemic was known previously from three localities in the Wrangell-St. Elias Mountains: Russell Glacier (61°42.0' N 141°45.0’ W, D. F. Murray 2170, 11 August 1968 (ALA)), Sheep Glacier (61°42.0’ N 141°39.0’ W, D.F. Murray 2251, 15 August 1968 (ALA)) and Chitistone Pass (61°37’ N 142°3’ W, R. Scott 1761 (MICH), Scott (1968) and from a station near Mentasta Pass (Hultén 1968). The specimens cited above extend the range 139 km west in the Wrangell Mountains, 85 km north into the Nutzotin Mountains and connect the distribution to the north- west in the Alaska Range with collections 40 km to the east in the Yukon Territory Cody (1996). It is rare in Alaska (GS S3) and in the Yukon Ternitory (Douglas et al. 1981). Map 94. Stellaria umbellata Turcz., Umbrella Starwort — CHUGACH MOUNTAINS: occasional in mesic tundra in limestone unit between Iron & Lime Creeks, 1798 m, 61°1’ N 141°53.18’ W, M. Cook 94430A, 8 August 1994; scattered in mud seeps, Verde Ridge, 1554 m, 61°14.03’ N 143°28.52’ W, Roland & D’Auria 96-478, 11 July 1996; occasional in moist calcareous gravel, Granite Creek, 1829 m, 61°0.22’ N 141°51.1’ W, C. Roland 96-737, 28 July 1996; moist N-facing scree, West Fork Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96- 312A, 29 July 1996 (ALA). NUTZOTIN MOUNTAINS: patchy in sheep bed at base of talus slope, Stone Creek, 1585 m, 62°25.78' N 142°22.11’ W, M. Cook 95101, 27 June 1995. St. ELIAS MOUNTAINS: rare in wet gravel, Mt. Chitina, 2073 m, 60°57.74' N 141°17.33'’ W, C. Roland 95-194, 14 July 1995; M.Cook 95248, 16 July 1995. WRANGELL MOUNTAINS: common in saturated soil of seep, Hasen Creek, 1835 m, 61°34.06' N 142°18.33' W, C. Roland 96-638, 23 July 1996. This amphiberingean arctic-alpine species is rare in Alaska (G4 §2S3) and the Yukon Territory (Douglas et al. 1981). It was known from Chitistone Pass in the Wrangell Mountains (61°37' N 141°58' W, D. F. Murray 1044, (CAN) (Murray 1968)). The specimens cited above extend its range 90 km south into the Chugach Mountains, 93 km north into the Nutzotin Mountains and connect its distribu- tion 125 km to the west in the Valdez Quad (Valdez Glacier, 61°08.0' N 146°10.0' W, L. A. & E.G. Viereck 2229, 10 August 1957 (ALA)) with a station 151 km to the east in the Yukon Territory (Cody 1996). Map 95. mr i me rn rr re eee | 244 THE CANADIAN FIELD-NATURALIST CERATOPHYLLACEAE Ceratophyllum demersum L., Coon’s Tail — COPPER RIVER BASIN: dominant, submerged in shallow water next to shore, pond at 2 km, Old Edgerton Highway, 415 m, 61°49.3’ N 145°10.7' W, M. Cook 95371, 8 August 1995. The specimen cited above extends the range of this species 201 km south from the Healy Quad (vic. Cantwell, 63°24.0' N 148°56.0' W, G. Smith 2243, 6 September 1953 (ALA)). It is rare in the Yukon Territory (Douglas et al. 1981). Map 96. NYMPHAEACEAE Nymphaea tetragona Georgi ssp. leibergii (Morong) Pors., Dwarf Water Lily — CHITINA RIVER BASIN: beaver pond east of Chititu Creek, 457 m, 61°22.16’ N 142°40.29' W, M. Duffy 91048, 8 July 1991; flow- ering along margins of pond 11 mi E of Chitina on the McCarthy Road, 427 m, 61°31.38’ N 144°10.14' W, M. Cook 95376, 8 August 1995. The specimens cited above extend the range of this species 174 km south from the Tanana River Basin (Scottie Creek, 62°41.0’ N 141°05.0’ W, L. Vining 35, 22 July 1979 (ALA)) and 282 km east from the Matanuska River Basin (E. LePage 20715, 1947 (S) (Hultén 1949)). Map 97. RANUNCULACEAE Caltha leptosepala DC., Mountain Marsh-Marigold — CHUGACH MOUNTAINS: mesic forb tundra, upper Tebay Lake, 1219 m, 61°12.18' N 144°23.8' W, M. Duffy 91112, 12 August 1991; forming continuous mats in wet moss stringers, West Fork Goat Creek, 1487 m, 60°59.8’ N 142°11.8’ W, M. Cook 96657, 31 July 1996; patchy in moist hummocky tundra in moist soil, Grant Creek plateau, 1250 m, 61°17.65’ N 143°56.34' W, Cook & Losso 96347, 7 July 1996; numerous in wet moss along creek, alpine valley 5.25 mi SE of Spirit Mountain, 1006 m, 61°16.84’ N 144°29.86' W, M. Cook 96444, 7 July 1996; occa- sional in mesic forb herbaceous meadow, vic. 12- mile Creek, 1326 m, 60°50.21’ N 142°30.85' W, M. Cook 96592, 7 July 1996; rare in moist areas around alpine lake, Nerelna Creek plateau, 1372 m, 61°26.66' N 144°17.69' W, Roland & D’Auria 96- 418, 8 July 1996; scattered in subarctic lowland sedge wet meadow, upper Tebay Lake, 579 m, 61°11’ N 144°24' W, Parker & Gracz 6751, 7 August 1996. St. ELIAS MOUNTAINS: alpine herba- ceous meadow, Karr Hills, 549 m, 60°8.66' N 141°16.44' W, K.A. Beck s.n., 12 August 1987. This species is North American with a cordilleran distri- bution. The collections cited above extend its range 192 km east into the Chugach Mountains and 267 km southeast into the southern St. Elias Mountains from a station at Thompson Pass (J.P. Anderson 1901, 30 July 1935 (S) (Hultén 1943)) and connect the range 195 km to the east in the Yukon Territory (Cody 1996). Map 98. Coptis trifolia (L.) Salisb., Trifoliate Goldthread — CHUGACH MOUNTAINS: ericaceous shrub bog, Middle Fork of the Bremner River, 869 m, 60°55.05’ N 143°43.86' W, Duffy & Barnes 96-096, 8 August Vol. 116 1996. GULF OF ALASKA: ericaceous shrub/graminoid herbaceous bog, between Porcupine Creek and Yakataga River, Robinson Mountains, 488 m, 60°4.8’ N 142°12.8' W, M. Duffy 9217, 11 June 1992. This species is amphiberingean with a boreal-montane distribution. The specimens cited above extend its range 91 km northeast in the Chugach Mountains from a collection at Katella (60°12.0' N 144°31.0’ W, J. Ver Hoef 29, 21 June 1979 (ALA)), 136 km east to the Robinson Mountains from a collection at Cape St. Elias (59°56.0’ N 144°23.0’ W, Cunningham & Stanford 130-78, 4 June 1978 (ALA)) and connect with a station 155 km to the east at Yakutat (F. Funston s.n., 1 July 1892 (US) (Coville and Funston 1896, Hultén 1941-1950)). Map 99. Delphinium brachycentrum Ledeb., Northern Larkspur — NUTZOTIN MOUNTAINS: scattered on extensive SW-facing gravel slopes, Baultoff Creek, 1707 m, 62°9.13' N 141°14.51' W, C. Roland 94- 160, 28 June 1994; occasional on unstable scree slope, headwaters of Alder Creek, 1554 m, 62°28.44’ N 142°15.06' W, M. Cook 95134, 30 June 1995. This amphiberingian arctic-alpine species is rare in the Yukon Territory (Douglas et al. 1981). The specimens cited above extend its range 203 km south into the Nutzotin Mountains from the Mt. Hayes Quad (Horn Mt, 63°38.0' N 144°44.0' W, L.A. Spetzman 154, 25 July 1957 (ALA)). Map 100. Ranunculus gelidus Karel. & Kiril. ssp. grayi (Britt.) Hult. — CHUGACH MOUNTAINS: occasional on W- facing scree slope, Iron Creek, 1798 m, 61°1’ N 141°53.18' W, M. Cook 94437, 6 August 1994; loose marble gravel on SW-facing slope, Granite Creek, 1829 m, 61°0.22' N 141°51.1’ W, C. Roland 96-745, 28 July 1996. MENTASTA MOUNTAINS: occa- sional in unstable scree, Devil’s Mountain, 1530 m, 62°25.62’ N 142°53.93' W, M. Cook 96280A, 7 July 1996; common in calcareous gravel, Totschunda Creek, 1280 m, 62°27.63' N 142°12.44' W, C. Roland 96-292, 24 June 1996; scattered on moist scree, headwaters of Lost Creek, 1722 m, 62°36.45' N 143°12.03' W, M. Cook 95166, 4 July 1995; upper Trail Creek, 1341 m, 62°37.09’ N 143°16.06’ W, M. Potkin 95-055, 28 July 1995; occasional in steep gully, Lost Creek, 1189 m, 62°35’ N 143°9.1’ W, Roland & D’Auria 97-061, 26 June 1997. NUTZOTIN MOuNTAINS: few on basalt tuff slope, Wiki Creek, _ 1411 m, 61°54.46’ N 141°10.68’ W, M. Cook 96088, 10 June 1996; S-facing talus slope, Rocker Creek, 1433 m, 61°54.8’ N 141°2.03’ W, M. Cook 94136, 23 June 1994; Carl Creek, 1920 m, 62°3.52’ N 141°36.27' W, M. Cook 95028, 16 June 1994; scattered on unstable scree, Alder Creek, 1554 m, 62°28.44' N 142°15.06’ W, M. Cook 95133, 30 June 1995; rare in platy rubble on bluff, Rock Lake, 1119 m, 61°48.7’ N 141°16.57' W, C. Roland 96-036, 5 June 1996; rare on steep talus slope, Wiki Creek, 1411 m, 61°54.46' N 141°10.68' W, C. Roland 96- 075, 8 June 1996; SW-facing ravine, Baultoff Creek, 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 96. Ceratophyllum demersum 245 _——so a een Sl ll See” 246 Vol. 116 100. Delphinium brachycentrum 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 101. Ranunculus gelidus ssp. grayi 104. Ranunculus sulphureus var. sulphureus 247 ee ee ee 248 1707 m, 62°9.13’ N 141°14.51’' W, Cook & Roland 94-144B, 94189A, 27 June 1994; occasional in unsta- ble granite scree, Carl Creek, 1920 m, 62°3.52' N 141°36.27' W, M. Cook 95009, 14 June 1994; Klein Creek, 1747 m, 62°2.29’ N 141°19.88’ W, M. Cook 95011, 15 June 1995; volcanic plateau between Bryan and Willow Creeks, 1829 m, 61°58.97' N 141°50.03’ W, M. Cook 95050, 18 June 1995. WRANGELL MOUNTAINS: Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24’ W, Moran & Roland 95-43, | July 1995; Lime Creek, 1646 m, 61°46.25’ N 141°50.36’ W, M. Cook 94114, 21 June 1994; Nabesna Glacier moraine, 1768 m, 61°56.6' N 143°0.00 W, M. Cook 9266, M. Duffy 92114, 26 June 1992: Lime Creek, 1707 m, 61°28.41’ N 141°29.44’ W, M. Cook 92497, M. Duffy 92143, 7 July 1992; Alice Peak, 1628 m, 61°39.82’ N 144°7.76’ W, C. Roland 96-392, 5 July 1996; Iron Mountain, 1868 m, 61°37.85' N 144°1.01' W, C. Roland 96-584, 16 July 1996; Chitistone Falls, 1177 m, 61°32.27’ N 142°11.49' W, C. Roland 96-628, 22 July 1996; Fish Creek, 1067 m, 62°16.92’ N 142°58.99’ W, M. Cook 95319, 27 July 1995; Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5’ W, M. Potkin 95-123, 30 July 1995; Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85’ W, M. Cook 95275, 18 July 1995; north slope of Mt. Santora. 722 ta, 62°23.4" N 144°15.9" WG Roland 94-006, 6 June 1994; Cone Ridge, 2073 m, 62°8.21’ N 143°18.47'’ W, C. Roland 94-305B, 27 July 1994 (WRST); Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24’ W, C. Roland 95-111, 1 July 1995. This species is amphiberingean with a cordilleran distri- bution. It was collected in the St. Elias Mountaints at Russell Glacier (61°42.0’ N 141°45.0’ W, D.F. Murray 2190, 11 August 1968 (ALA)) and Guerin Glacier (61°37.0’ N 141°50.0' W, D.F. Murray 2041, 3 August 1968 (ALA)). The new localites cited above extend its range 188 km west into the Wrangell Mountains, 113 km north into the Nutzotin Mountains and 81 km south into the Chugach Mountains. Map 101. Ranunculus pacificus (Hult.) Benson, Pacific Buttercup — GULF OF ALASKA: herbaceous/alder scrub, Cape Yakataga, 50 m, 60°3.32’ N 142°18.3’ W, M. Duffy 9205, 10 June 1992. The specimen cited above from Cape Yakataga connects the distribution of this Pacific Coastal species 418 km to the west in the Tyonek Quad (Lower Fish Creek, 61°22.0' N 150°31.0' W, J.A. Erickson 13, 18 June 1963 (ALA)) with collections 160 km to the southeast near Yakutat (Ankow River, E.P. Walker s.n., 1904 (US) and L.V. Piper 4468, 1904 (US) (Hultén 1943)). It is rare in Alaska (G3 S3). Map 102. Ranunculus pedatifidus Sm. ssp. affinis (R. Br.) Hult., Northern Buttercup — MENTASTA MOUNTAINS: mesic fissure of outcrop, Lost Creek, 1646 m, 62°34.58’ N 143°5.58'’ W, C. Roland 94-261, 23 July 1994. NUTZOTIN MOUNTAINS: Dryas tundra, Beaver Butte, 1829 m, 61°58.97’ N 141°50.03’ W, M. Cook 95054, 18 June 1995; locally abundant in turfy area, Carl THE CANADIAN FIELD-NATURALIST Vol. 116 Creek, 1920 m, 62°3.52' N 141°36.27’ W, C. Roland 95-023, 13 June 1994. St. EL1As MOUNTAINS: mesic meadow in scrub, White River, 1158 m, 61°44.87' N 141°45.4’ W, M. Duffy 92139, 6 July 1992. WRANGELL MOUNTAINS: few in rocky area, Nabesna River bluff, 1106 m, 62°15.05’ N 142°54.68' W, M. Cook 96185, C. Roland 96-163, 19 June 1996. This species is circumpolar with an arctic-alpine distri- bution. These localites extend its range south 126 km into the Wrangell Mountains and 144 km into the Nutzotin Mountains from a station near Northway (Hultén 1968). Map 103. Ranunculus sulphureus Soland, var. sulphureus Sulphur Buttercup —- NUTZOTIN MOUNTAINS: moist sedge meadow, Solo Flats, 1402 m, 62°11’ N 143°31.42’ W, M. Duffy 92155, 7 July 1992; occa- sional in seeps, Wiki Creek, 1411 m, 61°54.46’ N 141°10.68’ W, C. Roland 96-065, 8 June 1996; abundant in wet seeps, Beaver Butte, 1829 m, 61°58.97' N 141°50.03’ W, C. Roland 95-058, 18 June 1995. WRANGELL MOUNTAINS: Jaegar Mesa, 1893 m, 62°15.9' N 143°1.24’ W, Moran & Roland 95-29, 30 June 1995; mesic sedge meadow on tun- dra, Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5’ W, Potkin & Leggett 95-086, 29 July 1995; sedge tun- dra, Cheshnina Plateau, 1640 m, 61°50:33' N 144°22.5' W, M. Duffy 91089, 21 July 1991. This circumpolar arctic-alpine plant was collected at Skolai Pass in the Wrangell Mountains (Scott 1968). The specimens cited above extend its range 149 km east into the Wrangell Mountains. Map 104. Ranunculus trichophyllus Chaix in Villars var. erad- icatus (Laestadius) W.B. Drew, Thread-Leaf Water- Crowfoot — CHUGACH MOUNTAINS: Grant Creek, 853 m, 61°18.16' N 143°51.87' W, Cook & Losso 96337, 7 July 1996. WRANGELL MOUNTAINS: occa- sional in shallow water of pond, 911 m, 62°24.53’ N 143°42.68’ W, C. Roland 96-918, 11 August 1996; floating aquatic, Fish Creek, 1067 m, 62°16.92’ N 142°58.99’ W, M. Cook 95324A, 29 July 1995. This species is cirumpolar with a widespread distribu- tion. The specimens cited above extend its range south 181 km into the Wrangell Mountains and 301 km into the Chugach Mountains from a station near Healy Lake (Hultén 1968). Map 105. PAPAVERACEAE Papaver alboroseum Hult., Pale Poppy — CHUGACH MOUNTAINS: talus slope, Flood Creek, 1829 m, 61°42.88' N 141°58.65' W, M. Duffy 91059, 16 July 1991; rare on S-facing boulder slope, Juniper Island, 1291 m, 60°36.24' N 142°21.69’ W, M. Cook 94259, 11 July 1994; rare on W-facing limestone scree, ridge between Iron & Lime Creeks, 1798 m, 61°1’ N 141°53.18’ W, M. Cook 94439, 8 August 1994; occasional in old slide debris, pass between Amy Creek and Klu River, 1164 m, 61°4.01' N 143°34.83'’ W, C. Roland 96-530, 13 July 1996; few in loose rubble and sand, E Fork Kiagna River, 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 249 1487 m, 60°59.8’ N 142°11.8' W, C. Roland 96-844, 31 July 1996. NuTzoTIn MounrTaAINs: rare on S-fac- ing talus slope, Carl Creek, 1920 m, 62°3.52’ N 141°36.27' W, M. Cook 94112A, 20 June 1994; rare on barren S facing granidorite slope, Carl Creek, 1859 m, 62°3.07' N 141°35.01’ W, M. Cook 94128, 22 June 1994. St. ELIAS MOUNTAINS: scattered on scree slopes, Mt. Chitina, 2073 m, 60°57.74’ N 141°17.33’ W, M. Cook 95237, C. Roland 95-198B, 95-208, 15 July 1995. WRANGELL MOUNTAINS: rare in granitic scree, Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85’ W, C. Roland 95-241, 18 July 1995; few in gravel scree, Iron Mountain, 1868 m, 61°37.85'’ N 144°1.01’ W, M. Cook 96464, 16 July 1996; rubble and talus slope, Bonanza Ridge, 1768 m, 61°30.7’ N 142°49.85’ W, G. Dodge s.n., 10 August 1993; patchy on barren slope, Grotto Creek, 1847 m, 61°30.56’ N 142°24.79' W, M. Cook 96502, 23 July 1996; few in gravel scree, alpine lake basin E side of Lakina Glacier, 1219 m, 61°33.64’ N 143°19.01' W, M. Cook 96548, 24 July 1996; gently sloping gravel barrens, Nikolai Mine, 1695 m, 61°27' N 142°39’' W, Batten & Barker 96-081, 24 July 1996; rare in moist gravel, volcanic ridge 1 km east of Snyder Peak, 1524 m, 62°4.47’ N 144°30.51’ W, C. Roland 96-384, 5 July 1996; few in moist gravel, Cheshnina Plateau, 1399 m, 61°48.04’ N 144°6.27' W, M. Cook 96711, 5 August 1996. This poppy, rare in Alaska (G3G4 S3), was previously known from two localities in the Wrangell-St. Elias Mountains: Sheep Glacier (61°42.48’N 141°38.59'W, D. F. Murray 2212, 14 August 1968 (ALA) (Murray 1971)) and Bonanza Ridge (61°30’ N 142°51' W, Nordell & Schmitt 1, 101, 160, 166, 171, 301, 464 (LD & ALA) (Nordell & Schmitt)). It is now known from 17 additional localities in the Park including the Nutzotin and Chugach Mountains up to 108 km distant from the earlier collections. These collec- tions are 161 km disjunct from the range of this species to the west in the Anchorage Quad (Eklutna Glacier, 61°17.0' N 148°58.0' W, S.L. Welsh 4178, 18 June 1965 (ALA)) and 135 km south of the range to the north in the Mt. Hayes Quad (Rainbow Ridge, 63°15.0’ N 145°40.0’ W, C. Parker 1710, 2 August 1985 (ALA)). Map 106. Papaver radicatum Rottball ssp. kluanense (D. Love) D.F. Murray (P. kluanensis D. Love, P. freedmani- anum D. Love), Arctic Poppy — MENTASTA MOUNTAINS: rare in runnels of soil in fine limestone scree with alpine herbs, ridge between Little Jack and Trail Creeks, 1615 m, 62°36.05' N 143°17.63' W, C. Roland 95-010, 6 June 1995. NuTZOTIN MOUNTAINS: few on rhyolite talus slope with sparse Dryas-lichen- alpine herbs, vic. Wiki Peak, 1433 m, 61°53.9' N 141°9.51' W, M. Cook 94126, 16 June 1994; few in unstable, granodiorite scree, S-facing slope with sparse alpine herbs, Carl Creek, 1859 m, 62°3.07' N 141°35.01’ W, M. Cook 94127A, 22 June 1994; few localized on barren, unstable, granodiorite cobble and boulder scree S-facing slope, 1585 m, 62°2.87' N 141°34.46' W, M. Cook 95002, C. Roland 95-029, 13 June 1995; occasional in crevices and in gravel scree of boulder field, sparsely vegetated with sagebrush- herbs, Rock Lake, 1119 m, 61°48.7’ N 141°16.57' W, M. Cook 96029, 4 June 1996; few scattered on unsta- ble steep S-facing rhyolite scree knob, Wiki Basin, 1524 m, 61°54.77' N 141°11.05’ W, M. Cook 3186, 15 July 1998. St. ELIAS MOUNTAINS: few in exposed ash, sparsely vegetated with seral herbs, Cub Creek, 1280 m, 61°37.27' N 141°9.75' W, M. Cook 94185, 16 June 1994; scattered at base of limestone knob in ash with seral herbs and Dryas, Mt. Natazhat, 1716 m, 61°35.38’ N 141°1.83' W, M. Cook 95070, 95078, C. Roland 95-064, 95-072, 16 June 1995; occasional on barren N-facing gravel slope, Mt. Chitina, 2073 m, 60°57.74’ N 141°17.33’ W, C. Roland 95-221, 16 July 1995. WRANGELL MOUNTAINS: scat- tered in frost-boils on high elevation ridge of lime- stone, Cooper Pass, 1942 m, 62°17.16' N 142°31.44’ W, C. Roland 94-278, 24 July 1994; scattered in bare mineral soil on S-facing slope, volcanic ridge between Jacksina River & Mesa Creek, 2073 m, 62°8.21' N 143°18.47’ W, M. Cook 94374, 25 July 1994; large population in volcanic tuff, ridge between upper Jacksina River & Mesa Creek, 2073 m, 62°8.21' N 143°18.47' W, M. Cook 94407, C. Roland 94-312, 94- 313, 26 July 1994. This North American cordilleran poppy was known from the St. Elias Mountains of Alaska and the Yukon, vicinity of Mayo Lake in the Yukon and Eklutna Valley near Anchorage. The new localities cited above extend the range of this subspecies into the Mentasta, Nutzotin and Wrangell Mountains as far as 126 km from the earlier col- lections. Except for the locality disjunct 338 km to the west in the Anchorage Quad (Eklutna Valley, 61°26.0’ N 149°08.0’ W, L. C. Marvin 2114, 19 Jul 1985 (ALA)), these collections represent the western extension of the range of this subspecies. Map 107. Papaver walpolei A.E. Porsild, Walpole’s Poppy — MENTASTA MOUNTAINS: scattered on rubble slope, headwaters of Lost Creek, 1722 m, 62°36.45' N 143°12.03’ W, M. Cook 95172, 4 July 1995. This poppy is amphiberingean with an arctic-alpine dis- tribution. It is rare in Alaska (G3 S3) and in the Yukon Territory (Douglas et al. 1981). The locality cited above represents the southern limit of its distribution in Alaska and the Yukon, is 970 km disjunct from populations on the Seward Peninsula in northwestern Alaska (Solomon Quad: 64°34.0' N 163°43.0' W, S.D. Kildaw s.n., July 1987 (ALA)) and 325 km southwest of collections near Dawson in the Yukon Territory (Cody 1996). Map 108. BRASSICACEAE Aphragmus eschscholtzianus Andrz., Aleutian Cress — CHUGACH MOUNTAINS: moist sandy runnels on NE-facing stony slope, Verde Ridge, 1554 m, 61°14.03’ N 143°28.52' W, Roland & D’Auria 96- 471, 96-481, 10 July 1996; occasional in moist cal- careous gravel, Granite Creek, 1829 m, 61°0.22' N 141°51.1' W, C. Roland 96-739, 28 July 1996; moist bouldery solifluction slope, West Fork Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96- me ne ee ee Cie 250 305, 29 July 1996; scattered in granite gravel on ridge, East Fork Kiagna River, 1487 m, 60°59.8' N 142°11.8’ W, M. Cook 96652, 31 July 1996; few in wet mineral soil, Nelson Mountain, 1558 m, 61°19.37' N 143°48.83’ W, M. Cook 96371, 7 July 1996; few amongst unstable shale-gravel, Grant Creek plateau, 1250 m, 61°17.65’ N 143°56.34' W, M. Cook 96410, 7 July 1996; occasional in gravel stringers Bridge Creek plateau, 1871 m, 61°20.41’ N 144°6.97' W, M. Cook 96492, 7 July 1996; common in mesic tundra, Nerelna Creek plateau, 1372 m, 61°26.66' N 144°17.69' W, Roland & D’Auria 96- 402, 8 July 1996. MENTASTA MOUNTAINS: occasional in mesic alpine tundra, headwaters of Lost Creek, 1722 m, 62°36.45’ N 143°12.03' W, M. Cook 95161, 7 July 1995. NUTZOTIN MOUNTAINS: scattered in wet moss and bare organic soil, Cooper Pass, 1942 m, 62°17.16' N 142°31.44' W, M. Cook 94345, 24 July 1994; moist bare mineral soil on NE-facing boulder slope, Klein Creek, 1747 m, 62°2.29' N 141°19.88' W, M. Cook 95015, 15 June 1995; occasional in wet mossy seep, volcanic plateau between Bryan and Willow Creeks, 1829 m, 61°58.97’ N 141°50.03’ W, C. Roland 95-050, 18 June 1995; wet moss seep, Antler Creek, 1585 m, 62°25.78'’ N 142°22.11’ W, K.A. Beck 95-200, 27 June 1995. ST. ELIAS MOUNTAINS: rare in moist bare organic soil of solifluction lobes, limestone ridge between Dan & Copper Creeks, 1554 m, 61°21.56’ N 142°26.43’ W, M. Cook 94292, 14 July 1994; scattered in wet seep, Mt. Chitina, 2073 m, 60°57.74' N 141°17.33’ W, M. Cook 95225, 14 July 1995. WRANGELL MOUNTAINS: alpine basin, northwest flank of Mt. Drum, 1615 m, 62°4.5’ N 144°45.91’ W, C.R. Meyers 84-29, 4 July 1984; sedge tundra frost boil, Cheshnina Plateau, 1640 m, 61°50.33’ N 144°22.5’ W, M. Duffy 91088, 21 July 1991; common in mesic situations, Cone Ridge, 2073 m, 62°8.21’ N 143°18.47’ W, C. Roland 94-306, M. Cook 94362, 25 July 1994; occasional in bare sandy gravel, Chetaslina Plateau, 1615 m, 61°56.51' N 144°25.93'’ W, M. Cook 94475, 15 August 1994; common in stony streambed, Jaegar Mesa, 1893 m, 62°15.9' N 143°1.24’ W, C. Roland 95-107, 30 June 1995; patchy in wet moss, Crystalline Hills, 1585 m, 61°23.59' N 143°31.85’ W, M. Cook 95263, 17 July 1995, C. Roland 95-243, 18 July 1995; alluvial fan of seep, streambed & sorted rocks, Lakes Plateau, 1890 m, 62°4.4' N 143°23.5’ W, A. Leggett 95-092, 95-091, 95-079, 95-085, 95-103, 28 & 30 July 1995; scattered in loose, moist limestone gravel on N-facing slopes, Iron Mountain, 1868 m, 61°37.85’ N 144°1.01' W, C. Roland 96-578, M. Cook 96455, 16 July 1996; scattered in clay soil on east facing slope, Grotto Creek, 1847 m, 61°30.56’ N 142°24.79’ W, C. Roland 96-649A, 23 July 1996; moist muddy scree, Nikolai Mine, 1695 m, 61°27’ N 142°39’ W, Batten & Barker 96-095, 27 July 1996 (ALA); rare in snowbed areas, upper Dadina River, 1646 m, 62°3.88' N 144°35.76' W, C. Roland 96-371, 5 July 1996; THE CANADIAN FIELD-NATURALIST Vol. 116 occasional in protected chute, limestone ridge south- west of Alice Peak, 1628 m, 61°39.82’ N 144°7.76' W, C. Roland 96-388, 5 July 1996; occasional in mesic volcanic gravel stringers, Hasen Creek, 1835 m, 61°34.06'’ N 142°18.33' W, M. Cook 96494, 7 July 1996; occasional in moss on limestone outcrops, Lakina Glacier, 1219 m, 61°33.64’ N 143°19.01' W, M. Cook 96538, 7 July 1996; moist silt adjacent to creek, Nikolai Pass, 1280 m, 61°26’ N 142°40’ W, Batten & Barker 96-045, 7 July 1996 (ALA). This Alaska-Yukon endemic with a Pacific Coastal dis- tribution was known previously from only two localities in the Park: Chitistone Pass (61°37’ N 141°58’ W, D. F. Murray 1041 (CAN) (Murray 1968)) and Bonanza Ridge (61°30' N 142°51’ W, Nordell & Schmitt 163 (LD & ALA) (Nordell and Schmitt 1978)). As indicated by the above collections, this species is now known from throughout the mountain ranges in the Park where it is locally common in alpine mesic sites. Collections from the western Wrangells and the Mentasta Mountains are up to 150 km distant from prior Park collections, and the closest station to the speci- mens cited above is 78 km to the southwest at Thompson Pass (Valdez Quad: 61°08.0’ N 145°45.0’ W, C.L. Parker 2415, 22 July 1990 (ALA)). Map 109. Arabis calderi G. A. Mulligan (Arabis lyallii sensu Hultén (1968) pro parte), Calder’s Rockcress — ST. ELIAS MOUNTAINS: scattered in meadow adjacent to sheep trail and in open soil on S-facing slopes, ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94' N 141°6.75’ W, C. Roland 95-169, M. Cook 95215, 12 & 13 July 1995. This North American cordilleran species is new to the flora of Alaska. The western limit of its distribution is rep- resented by the above collection, the closest station being 172 km to the east in the Yukon Territory (Cody 1996). Its distribution to the south and east of the Yukon Territory is spotty, ranging from Great Bear Lake of the Mackenzie District in the Northwest Territories (1108 km distant), to the southwest corner of Alberta and British Columbia (1654 km distant), south to Washington, California and Montana (Mulligan 1995; Cody 1996). It is rare in Alaska (G3G4 S1) and Cody et al. (1998) suggest that it be added to the list of rare species in the Yukon Territory. Map 110. Arabis codyi G.A. Mulligan, Cody’s Rockcress — CHUGACH MOUNTAINS: rare on W-facing unstable limestone scree, Granite Range, 1798 m, 61°1' N 141°53.18’ W, M. Cook 94441, 5 August 1994. This Rockcress, which is new to Alaska, was known from only two localities prior to our collection in the Chugach Mountains: the holotype 230 km to the east in the St. Elias Mountains, Yukon Territory (D.F. & B. Murray 72, 1 July 1965 (DAO)) and a specimen from British Columbia 1174 km to the southeast (Perow, 54 30’N, 126 26'W, Taylor & Levis 468 (UBC) (Mulligan 1995)). This species is considered rare in Alaska (G1G2 S1) and should be added to the list of rare species in the Yukon Territory (Cody et al. 1998). Map 111. Arabis drepanoloba Greene (Arabis lemmonii S. Wats. var. drepanoloba (E.L. Greene) Rollins), Rockcress —- CHUGACH MOUNTAINS: few in gravel scree between limestone outcrops, ridge above 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 108. Papaver walpolei 251 252 THE CANADIAN FIELD-NATURALIST Vol. 116 III. Arabis codyi 112. Arabis drepanoloba 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 116. Braya glabella ssp. purpurascens 253 - ----OOoOoOC Ooo Oe COC Sh 254 Canyon Creek, 1682 m, 61°24.28’ N 144°21.61’ W, Cook & Losso 96325, 7 July 1996 (ALA). This North American cordilleran Rockcress is also new to the flora of Alaska. The locality cited above, the western limit of its distribution, is 290 km west of a station in the Yukon Territory (Cody 1996). These populations are dis- junct from the main range in southeast British Columbia, Alberta, Colorado, Wyoming and Montana (Cody 1996, Mulligan 1995). It is rare in Alaska (GS $1) and should be added to the list of rare species of the Yukon Territory (Cody 1996). Map 112. Arabis lemmonii S. Wats., Lemmon’s Rockcress — CHUGACH MOUNTAINS: mesic shrub birch-ericaceous scrub, Granite Creek, Granite Range, 701 m, 60°43.78' N 142°31.4’ W, Duffy & Barnes 96-064, 8 August 1996 (ALA), confirmed by G. A. Mulligan (DAO) 1998. Another North America cordilleran Rockcress new to the flora of Alaska and with the western limit of its range in the Park. This species is also rare in Alaska (G5 S1) and in the Yukon Territory (Douglas et al. 1981). The specimen cited above is 116 km west of the Yukon Territory collec- tion near Kluane Lake (Cody 1996). The stations in Alaska and the Yukon are disjunct from the main range of this species in southwestern Alberta, southern British Columbia, Montana, Idaho, Washington, Wyoming, Colorado, Utah, Nevada, Oregon and California (Mulligan 1995). Map 113. Arabis media N. Busch (Arabis arenicola (Richardson ex Hook.) Galert var. pubescens S. Wats. pro parte), Rockcress — WRANGELL MOUNTAINS: scattered in loose gravel and sparse tun- dra on SW-facing slope, vic. Snyder Peak, 1524 m, 62°4.47' N 144°30.51' W, C. Roland 96-383, 5 July 1996; occasional in moist sandy soil, vic. Long Glacier, 1399 m, 61°48.04' N 144°6.27' W, M. Cook 96709, 8 August 1996 (ALA); scattered in Rhacomitrium moss on gravel bar, upper Copper River, 945 m, 62°19.02' N 143°47.44' W, M. Cook 96719, 8 August 1996. This amphiberingean arctic-alpine Rockcress was known previously from the vicinity of Guerin Glacier in the St. Elias Mountains (D. F. Murray 2022, 3 August 1968 (ALA)). The stations cited above extend the range 156 km west into the Wrangell Mountains. These southcentral Alaska collections are 913 km southeast of stations in west- ern Alaska and 821 km south of stations along the arctic coast of the Yukon Territory (Cody 1996, Hultén 1968). This species should be added to the list of rare plants of the Yukon Territory (Cody 1996). Map 114. Braya glabella Richardson ssp. glabella, Braya, Smooth Northern Rockcress — MENTASTA MOUNTAINS: rare in moist bare soil on limestone ledge above Soda Lake, 1173 m, 62°32.36' N. 142°53.98' W, M. Cook 94309, 21 July 1994; rare in limestone gravels, ridge between Soda and Platinum Creeks, 1494 m, 62°31.2' N 142°54.03’ W, C. Roland 95-134, 5 July 1995; rare in red colored cal- careous gravel on ridge above limestone outcrops, Baultoff Creek, 1707 m, 62°9.13’ N 141°14.51’ W, THE CANADIAN FIELD-NATURALIST Vol. 116 M. Cook 94781,°27 June 1994-" ST ees MOUNTAINS: barren scree slope, ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94’ N 141°6.75’ W, M. Cook 95205, 11 July 1995. This North American arctic-alpine mustard is rare in the Yukon Territory (Douglas et al. 1981). The specimens cited above extend its range 310 km south from Charley River Quad (Upper Hard Luck and Cathedral Creek, 65°09.0'’ N 141°03.0' W, Parker & Hasselbach 6340, 23 June 1996 (ALA)) and connect the range 54 km to the east in the Yukon Territory (Cody 1996). Map 115. Braya glabella Richardson ssp. purpurascens (R. Br.) Cody (Braya purpurascens (R. Br.) Bunge), Purplish Braya — CHUGACH MOUNTAINS: rare on W-facing limestone scree slope & in Dryas dwarf scrub tundra, ridge between Iron and Lime Creeks, 1798 m, 61°1’ N 141°53.18’ W, M. Cook 94435, 94445B, 6 August 1994. WRANGELL MOUNTAINS: rare on boulder slope, Rambler Mine, 1219 m, 62°21.92’ N 143°2.87' W, M. Cook 8979, 9 June 1989. The southern limit of this circumpolar arctic-alpine mus- tard is documented by the above localities. These collec- tions extend the range 242 km southeast into the Chugach and Wrangell Mountains from the Alaska Range (Healy Quad: Dry Creek, 63°51.0’ N 147°30.0’ W, Viereck & Jones 5953, 23 July 1962 (ALA)) and connect the range 250 km to the east in the Yukon Territory (Cody 1996). This species is rare in the Yukon Territory (Douglas et al. 1981). Map 116. Draba cinerea Adams, Grayleaf Whitlowgrass — MENTASTA MOUNTAINS: barren, gravel scree slope on limestone ridge between Lost and Platinum Creeks, 1646 m, 62°34.58’ N 143°5.58’ W, M. Cook 94329A, 23 July 1994. St. ELIAS MOUNTAINS: scattered on loose, disturbed, S-facing marble scree slope between Logan and Walsh Glaciers, 1951 m, 60°53.94' N 141°6.75' W, C. Roland 95-148, 11 July 1995. WRANGELL MOUNTAINS: rare under boulder on basalt tuff and rubble slope, Cone Ridge, 2073 m, 62°8.21’ N 143°18.47’ W, M. Cook 94396, 25 July 1994. This species is circumpolar with an arctic-alpine distri- bution. It was collected adjacent to the Park in 1981 at the confluence of the Chitina and Copper Rivers (D. F. Murray 6194 (ALA)) which is 79 km south of the specimen cited above from Cone Ridge and 190 km west of the station in the St. Elias Mountains. These collections in southcentral Alaska are 804 km disjunct from the main range of the species in arctic Alaska (Demarcation Point Quad: Nuvagapak Pt., 69°53.0’ N 142°18.0’ W, C.R. Meyers 80-14, 20 July 1980 (ALA)) and 634 km northeast of a col- lection in the Mt. Katmai Quad (Coville Lake, 58°45.0' N 155°37.0’ W, G. Schaller s.n., 8 July 1954 (ALA)). Map 117. Draba corymbosa R. Br. ex DC. (D. macrocarpa Adams), Flattop Whitlowgrass — CHUGACH MOUNTAINS: occasional in mossy streamside tundra on SE-facing slope, Verde Ridge, 1554 m, 61°14.03’ N 143°28.52' W, Roland & D’Auria 96-476, 11 July 1996; occasional in moist calcareous gravel in pro- 2002 - COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 255 tected area on north side of ridge, Granite Creek, 1829 m, 61°0.22' N 141°51.1’ W, C. Roland 96- 741B, 28 July 1996; W-facing rocky tundra slope, East Fork Kiagna River, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-326C, 29 July 1996; occa- sional on barren limestone shale at summit of plateau, Bridge Creek, 1871 m, 61°20.41' N 144°6.97' W, M. Cook 96489, 7 July 1996. WRANGELL MOUNTAINS: rare in rocks on S-facing slope, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24’ W, C. Roland 95-117C, 1 July 1995; Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5' W, A. Leggett 95-083, 30 July 1995; moist clay gravel flats, Nikolai Mine, 1695 m, 61°27’ N 142°39' W, Batten & Barker 96-090, 24 July 1996 (ALA); steep W-SW-facing limestone scree, Chitistone Mountain, 1219 m, 61°28’ N 142°33' W, Batten & Barker 96- 149A, 26 July 1996 (ALA). This mustard is circumpolar with an arctic-alpine distri- bution. The stations cited above are 247 km southeast of stations in the Alaska Range (Hultén 1968) and 156 km west of stations in the Yukon Territory where it is widely distributed throughout the mountains (Cody 1996). Map 118. Draba crassifolia Graham, Snowbed Whitlowgrass — CHUGACH MOUNTAINS: Iron Creek, limestone unit between Iron & Lime Creeks, S of Chitina River, Granite Range, 1798 m, 61°1’ N 141°53.18' W, M. Cook 94433, 6 August 1994; boulder field on W-fac- ing slope, Goat Creek, 1487 m, 60°59.5' N 142°1' W, Batten & Barker 96-328A, 29 July 1996 (ALA); scat- tered in granite gravel on ridge, West Fork Goat Creek, 1487 m, 60°59.8’ N 142°11.8' W, M. Cook 96654B, 31 July 1996; few in bare mineral soil between gravel and cobble of rock outcrops, Nelson Mountain, 1558 m, 61°19.37' N 143°48.83’ W, M. Cook 96376, 7 July 1996. MENTASTA MOUNTAINS: idee east of Soda Lake; 1173 m, 62°32.36'’ N 142°53.98' W, C. Roland 94-241, 21 July 1994; unstable, nearly bare sandy area in limestone scree, Lost Creek, 1646 m, 62°34.58’ N 143°5.58' W, C. Roland 94-267, 23 July 1994. NUTZOTIN MOUNTAINS: exposed NW-facing talus slope, vic. peak 5745, Gold Hill, 1585 m, 62°4.05’ N 141°52’ W, M. Cook 92071, 27 June 1992; cutbank below sloughing Cassiope heath, Wiki Creek, 1411 m, 61°54.46’ N 141°10.68' W, M. Cook 96086, 10 June 1996. ST. ELIAS MOUNTAINS: rare in mesic, protected area on north side of small rock outcrop, ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94’ N 141°6.75' W, C. Roland 95-160, 12 July 1995; occasional in mixed soil and gravel of forb herbaceous community, Mt. Chitina, 2073 m, 60°57.74’ N 141°17.33’ W, C. Roland 95-196, 95-2015A, M. Cook 95240, 14 & 15 July 1995. WRANGELL MOUNTAINS: N and E-facing scree slope, Lime Creek ridge, 1935 m, 61°28.41' N 141°30.7’ W, M. Cook 92509A, 9 July 1992; uncom- mon, in heavily grazed sheep-bed area of turfy graminoid-forb tundra, Lime Creek ridge, 1646 m, 61°46.25’ N 141°50.36' W, C. Roland 94-102, 21 June 1994; open soil in tundra, Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85’ W, C. Roland 95-232B, 18 July 1995; occasional in moss of wet seep, Hasen Creek, 1835 m, 61°34.06’ N 142°18.33’ W, M. Cook 96497, 7 July 1996. NUTZOTIN MOUNTAINS: rockfield, ridge 2.8 km W of Ophir Creek, 1966 m, 61°56.1’ N 141°35.68' W, M. Duffy 92172, 8 July 1992. This amphiatlantic mustard with an arctic-alpine distri- bution was previously known from Skolai-Chitistone Pass in the Wrangell Mountains (61°37.0' N 141°58.0’ W, D. F. Murray 724, 1966 (ALA) & 61°37’ N 142°3' W, R. Scott s.n. (MICH)). It is now known throughout the alpine regions of the Park. The collections cited above connect its distribution 100 km to the northwest in the Alaska Range with its range 71 km to the east in the Yukon Territory (Cody 1996) where it was considered rare (Douglas et al. 1981). Map 119. Draba densifolia Nutt. ex Torrey & A. Gray, Denseleaf Whitlowgrass — NUTZOTIN MOUNTAINS: rockiieldy Ophir Créeck,.1966 m, 617°56.1' N 141°35.68' W, M. Duffy 92172, 8 July 1992. This North American cordilleran mustard is disjunct in Alaska from southern British Columiba, California, Oregon, Idaho, Montana, Wyoming and Utah. The collec- tion cited above from the Nutzotin Mountains is one of only seven localities in Alaska, the closest being 405 km to the northeast at Cantwell (A.E. & R.T. Porsild 61, 8 June 1926 (CAN, S) (Hultén 1941-1950)). It is rare in Alaska (G5 S1) and in the Yukon Territory where it is known in the Tombstone region of the Ogilvie Mountains (Cody 2000). Map 120. Draba incerta Payson, Yellowstone Whitlowgrass — CHUGACH MOUNTAINS: graminoid forb herba- ceous meadow, Iron Creek, 1798 m, 61°1’ N 141°53.18’ W, M. Cook 94442, 5 August 1994; rare in old slide debris, Amy Creek, 1164 m, 61°4.01' N 143°34.83' W, C. Roland 96-531, 13 July 1996; few in unstable limestone scree, Canyon Creek, 1682 m, 61°24.28’ N 144°21.61’ W, Cook & Losso 96326, 7 July 1996; few in bare mineral soil, Nelson Mountain, 1558 m, 61°19.37' N 143°48.83' W, M. Cook 96377, 7 July 1996; scattered in granitic rubble on S-facing slope, Nerelna Creek Plateau, 1372 m, 61°26.66' N 144°17.69' W, Roland & D’Auria 96- 410, 8 July 1996; rock outcrops, Towhead Mountain, 1433 m, 6193.21’ N 142°39.8' W, Duffy & Barnes 96-156, 8 August 1996. MENTASTA MOUNTAINS: Dryas-rock tundra, Lost Basin, 1722 m, 62°36.45' N 143°12.03’ W, M. Cook 95170, 7 July 1995. ST. ELIAS MOUNTAINS: rare in mesic area on small rock outcrop, ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94’ N 141°6.75'’ W, C. Roland 95- 159, 12 July 1995; rock crevices of S-facing outcrop, Mt. Chitina, 2073 m, 60°57.74' N 141°17.33' W, M. Cook 95235, C. Roland 95-201, 95-202, 95-206, 15 July 1995; N-facing scree, Nikolai Pass, 1280 m, 61°26’ N 142°40' W, Batten & Barker 96-016B, 23 July 1996. WRANGELL MOUNTAINS: scattered on bar- ren slope, Grotto Creek, 1847 m, 61°30.56' N 142°24.79' W, C. Roland 96-651, 23 July 1996. ee ee ee SS CS ee 256 THE CANADIAN FIELD-NATURALIST 120. Draba densifolia Vol. 116 2002 - COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 121. Draba incerta 124. Draba lonchocarpa var. thompsonii 257 Oe eee ee eC 258 This North American cordilleran mustard is rare in Alaska (G5 §2S3). The localities cited above connect its range 204 km to the west in the Anchorage Quad (Mi 119 Seward Highway, 61°07.0’ N 149°52.0' W, C. L. Parker 6953, 20 May 1997 (ALA)) with its distribution 88 km to the east in the Yukon Territory (Cody 1996). Map 121. Draba kananaskis G.A. Mulligan, Longstalk Whitlowgrass — CHUGACH MOUNTAINS: scattered in seeps on E-facing slope, Verde Ridge, 1554 m, 61°14.03’ N 143°28.52' W, Roland & D’Auria 96- 479, 11 July 1996; occasional in moist calcareous gravel, Granite Creek, 1829 m, 61°0.22’ N 141°51.1’ W, C. Roland 96-740, 96-741A, 28 July 1996. This North American cordilleran mustard was known only from the vicinity of Hope on the Kenai Peninsula (Hultén 1968). The two localities cited above from the Chugach Mountains are 318 km east of the Kenai Peninsula locality. Map 122. Draba lactea Adams — CHUGACH MOUNTAINS: rare on W-facing unstable scree slope, Iron Creek, 1798 m, 61°1’ N 141°53.18’ W, M. Cook 94445C, 6 August 1994. MENTASTA MOUNTAINS: scattered on N-facing limestone ledge in sheep bed, Lost Basin, 1722 m, 62°36.45' N 143°12.03’ W, M. Cook 95168, 4 July 1995; scattered over sparsely vegetated dry forb herbaceous rubble slope, Lost Basin, 1722 m, 62°36.45’ N 143°12.03’ W, M. Cook 95173, 4 July 1995. NUTZOTIN MOUNTAINS: few on S-facing red basalt rubble slope, Wiki Creek, 1411 m, 61°54.46’ N 141°10.68' W, M. Cook 96091, 10 June 1996. WRANGELL MOUNTAINS: polygon tundra, Lakes Plateau, 1890 m, 62°4.4' N 143°23.5’ W, M. Potkin 95-069, 29 July 1995; scattered in moist limestone, Iron Mountain, 1868 m, 61°37.85'’ N 144°1.01’ W, C. Roland 96-573, 96-576, 16 July 1996; N-facing scree, Nikolai Pass, 1280 m, 61°26’ N 142°40’ W, Batten & Barker 96-016C, 23 July 1996 (ALA); scattered in loose limestone cobbles on steep, SE- facing slope, Alice Peak, 1628 m, 61°39.82’ N 144°7.76' W, C. Roland 96-390, 96-391, 96-395, 5 July 1996. This circumpolar arctic-alpine mustard was known from one locality in the Wrangell Mountains (Chitistone Pass, 61°37' N 142°3’ W, R. Scott s.n., 1967 (MICH) (Scott 1968); 61°37’ N 141°58’ W, D. F. Murray 1052, 1053, 21 July 1967 (CAN) (Murray 1968)). The specimens cited above extend the range 114 km west in the Wrangell Mountains and 125 km north into the Mentasta Mountains. These collections connect the range 122 km north in the Alaska Range (Mt Hayes Quad: 63°28.0’ N 145°49.0' W, Batten et al. 78-67, 30 June 1978 (ALA)) with the distribu- tion 114 km to the east in the Yukon Territory (Cody 1996). Map123 . Draba lonchocarpa Rydb. var. thompsonii (C.L. | Hitche.) Rollins — MENTASTA MOUNTAINS: scattered at base of scree slope below limestone ridge, head- waters. of Lost. Creek, 1722-m,:62°36.45 08 143°12.03' W, M. Cook 95184, 11 July 1995. This specimen documents the northern and western limit of the range of this North American cordilleran mustard, THE CANADIAN FIELD-NATURALIST Vol. 116 the only known locality of this variety in Alaska. It is 1100 km disjunct from the mountains of British Columbia and western Washington (Rollins 1993 and Mulligan 1974). Map 124. Draba macounii O.E. Schulz — MENTASTA MOUNTAINS: rare in bare mineral soil on S-facing limestone talus slope, Soda Lake, 1173 m, 62°32.36’ N 142°53.98' W, M. Cook 94312A, 22 July 1994; rare in red thin-bedded limestone scree, ridge between Lost and Platinum Creeks at peak 5240, 1646 m, 62°34.58’ N 143°5.58’ W, M. Cook 9434 1B, 23 July 1994; rare in bare argillite gravels and rock crevices on windswept ridge, ridge between Little Jack and Trail Creeks, 1615 m, 62°3603 3) 143°17.63' W, C. Roland 95-021, 7 June 1995; sad- dle near contact of limestone and Nikolai Greenstone, ridge between Little Jack and Trail Creeks, 1615 m, 62°36.05’ N 143°17/63" Wo at Potkin 95-56B, 7 June 1995; upper Trail Creek, 1341 m, 62°37.09’ N 143°16.06' W, Potkin & Leggett 95-044, 27 July 1995; limestone talus, head- waters of Lost Creek, 1722. m; 62°3645 i 143°12.03’ W, K.A. Beck 95-221, 4 July 1995; scat- tered in gravel around outcrop on limestone ridge, between Soda and Platinum Creeks, 1494 m, 62°31.2' N 142°54.03’ W, C. Roland 95-145, 6 July 1995. NUTZOTIN MOUNTAINS: Dryas tundra, ridge between Fogenbera Pass and Shotgold Creek, 1783 m, 61°53.28’ N 141°37.1' W, M. Cook 92458, 8 June 1992. St. ELIAS MOUNTAINS: occasional in dwarf scrub/herbaceous meadow tundra, N-facing rock ledges and S-facing bare solifluction lobes, ridge between Dan & Copper Creeks, 1554 m, 61°21.56’ N 142°26.43' W, M. Cook 94286A, 13 July 1994; rare in bare moist organic soil on S-facing limestone outcrop, ridge between Dan & Copper Creeks, 1554 m, 61°21.56’ N 142°26.43’ W, M. Cook 94296A, 14 July 1994; occasional on vegetated stringer of limestone talus slope, ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94’ N 141°6.75’ W, M. Cook 95200, 11 July 1995; mixed soil and rubble in debris flow on W-facing slope, ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94’ N 141°6.75' W, C. Roland 95-153, 11 July 1995. WRANGELL MOUNTAINS: alpine tundra, 152 m above and E of Nabesna Glacier moraine, 1768 m, 61°56.6’ N 143° W, M. Cook 9267, 26 June 1992; scree slope, N and E-facing, ridge between Lime Creek and Solo Flats, 1935 m, 61°28.41’ N 141°30.7' W, M. Cook 92505, 92510, 8 July 1992; seep area in talus, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24' W, Moran & Roland 95-30, 30 June 1995; occasional in solifluction soil below ridge, Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85’ W, M. Cook 95268, 18 July 1995; occasional on bar- ren NE-facing slope of fine gravelly scree, Crystalline Hills, 1585 m, 61°23.59' N 143°31.85’ W, C. Roland 95-245B, 18 July 1995; Lakes Plateau, 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 128. Draba porsildii 2o9 Cer ee eee i 260 1890 m, 62°4.4’ N 143°23.5' W, M. Potkin 95-095A, 30 July 1995; scattered in barren limestone gravel on SW-facing slope, Iron Mountain, 1868 m, 61°37.85’ N 144°1.01’ W, C. Roland 96-580, 16 July 1996; scattered in barren gravel on S-facing ridge, ridge W of Grotto Creek headwaters, 1847 m, 61°30.56’ N 142°24.79' W, C. Roland 96-649B, 23 July 1996; occasional in moist volcanic gravel scree, Iron Mountain, 1868 m, 61°37.85’ N 144°1.01' W, M. Cook 96461, 7 July 1996; few on saddle in gravel and patchy alpine forb herbaceous vegetation, between headwaters of King and Pouch Creeks, 1926 m, 61°38.99’ N 144°3.51' W, M. Cook 96469, 7 July 1996; few in gravel scree on sparsely vegetat- ed NE-facing slope, alpine lake basin on limestone, E side of Lakina Glacier, 1219 m, 61°33.64’ N 143°19.01’ W, M. Cook 96547, 7 July 1996. This North American cordilleran mustard has a spotty distribution from Alaska south to Colorado and Montana. The specimens cited above from all of the mountains in the Park connect its range 227 km to the northwest in the Alaska Range (Healy Quad: Dry Creek, 63°52.0' N 147°30.0’ W, Viereck & Jones 5962, 23 July 1962 (ALA)) with the distribution 80 km to the southeast in the Yukon Territory (Cody 1996). Map 125. Draba oligosperma Hook., Fewseed Whitlowgrass — CHUGACH MOUNTAINS: occasional in marble rub- ble on steep and dry SE-facing slope, ridge between Canyon and Pass Creeks, 1463 m, 61°24.35’ N 144°20.8' W, Roland & D’Auria 96-436, 9 July 1996. St. ELIAS MOUNTAINS: common in dry gravels on S-facing slope with sagebrush-forb vegetation, ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94' N 141°6.75’ W, C. Roland 95-175A, 12 July 1995, M. Cook 95212, 95213, 13 July 1995. This North America cordilleran mustard was collected by H. M. Laing in 1925 on the upper Chitina River most likely in the vicinity of our collection from the St. Elias Mountains (Porsild 1939; Hultén 1941-1950). The other station in Alaska is our collection in the Chugach Mountains 182 km to the east. The stations on the upper Chitina River are 121 km west of the closest localities in the Yukon Territory and are disjunct from the main cordilleran range of the species in Colorado, California and Washington (Rollins 1993). Map 126. Draba palanderiana Kjellman., Palander’s Whitlow- Grass — NUTZOTIN MOUNTAINS: occasional on S- facing limestone outcrops, Baultoff Creek, 1707 m, 62°9.13' N 141°14.51' W, M. Cook 94180, 27 June 1994. WRANGELL MOUNTAINS: moist gravel on scree slope, in tuft of Erigeron purpuratus, 1585 m, 61°23.59’ N 143°31.85’ W, M. Cook 95267, 18 July 1995. This mustard is an Alaska-Yukon endemic with an arctic-alpine distribution and is rare in Alaska (G4G5 S4). The localities cited above are 159 km northeast of a station near Valdez (Porsild and Cody 1980) and 141 km south- west of a station in the Yukon Territory (Cody 1996). Map 127. THE CANADIAN FIELD-NATURALIST Vol. 116 - Draba porsildii G. A. Mulligan, Porsild’s Whitlowgrass —CHUGACH MOUNTAINS: scree, Goat Creek, Granite Range, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-327B, 29 July 1996 (ALA); few in granite gravel, ridge above W. Fork Goat Creek, 1487 m, 60°59.8’ N 142°11.8’ W, M. Cook 96661, 31 July 1996. MENTASTA MOUNTAINS: rare in bare mineral soil on S-facing talus slope, Soda Lake, 1173 m, 62°32.36'’ N 142°53.98' Wai Gam 94312B, 22 July 1994; occasional in gravel areas on limestone outcrop, Trail Creek, 1615 m, 62°36.05’ N 143°17.63' W, C. Roland 95-004, 95-015, 6 & 7 June 1995 (ALA); upper Trail Creek, 1341 m, 62°37.09' N 143°16.06’ W, M. Potkin 95-037A, 27 July 1995; scattered in limestone gravel, Soda Lake ridge, 1494 m, 62°31.2’ N 142°54.03’ W, C. Roland 95-144, 7 July 1995 (ALA). NUTZOTIN MOUNTAINS: few in bare mineral soil, ridge at headwaters of Alder Creek, 1554 m, 62°28.44’ N 142°15.06’ W, M. Cook 95131, 26 June 1997 (ALA). S7) Heras MOUNTAINS: rare in disturbed area, Mt. Chitina, 2073 m, 60°57.74' N 141°17.33’ W, C. Roland 95-203, 15 July 11995: The seven localities in the Mentasta, Nutzotin and Granite Ranges cited above are the only other stations in Alaska besides those in the Black River Quad 458 km to the north (66°35.0’ N 141°10.0’ W, R. Lipkin 91-46, 10 June 1991; 66°36.0’ N 141°11.0’ W, R. Lipkin 91-96, 11 June 1991; 66°36.0’ N 141°11.0' W, R. Lipkin 91-141, 14 June 1991; 66°32.0' N 141°04.0’ W, R. Lipkin 91-153, 15 June 1991 (ALA)). Our collections are 101 km west of sta- tions in the Yukon Territory (Cody 1996). This species is a North American cordilleran endemic known from Colorado north to the Northwest Territories, Yukon and Alaska where it is considered rare (Douglas et al. 1981). Map 128. Draba ruaxes Payson & H. (D. exalata Ekman, D. ventosa A. Gray var. ruaxes (Payson & H. St. John)), Rainier Whitlowgrass — CHUGACH MOUNTAINS: rare in gravel scree, serpentinized ultramafic unit, Granitic Creek ridge, 1768 m, 61°6.19’ N 142°55.03’ W, M. Cook 94220, 8 July 1994. MENTASTA MOUNTAINS: rare in moist bare soil on ledge between limestone gravel scree, Soda Lake, 1173 m, 62°32.36’ N 142°53.98' W, M. Cook 94311A, 21 July 1994; occa- sional on barren gravel scree, Lost Creek, 1646 m, 62°34.58’ N 143°5.58' W, M. Cook 94331, 23 July 1994; scattered in runnels of fine limestone scree near ridge, Trail Creek, 1615 m, 62°36.05’ N 143°17.63' W, C. Roland 95-009, 6 June 1995; occasional in moss betwen limestone rubble on N-facing slope, headwaters of Lost Creek, 1722 m, 62°36.45’ N 143°12.03' W, K.A. Beck 95-225, 95-231, 95-241, M. Cook 95175, 95185, 7 July 1995; common in fine, red calcareous gravel on E-facing slope above Totschunda Creek, 1280 m, 62°27.63’ N 142°12.44’ W, C. Roland 96-289, 24 June 1996; few in basalt scree, ridge N of Totschunda Creek mouth, 1494 m, 62°28.28' N 142°40.5’ W, Cook & Shea 96285, 7 July 1996; scattered in unconsolidated limestone and 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 261 greenstone rubble, Lost Creek, 1189 m, 62°35’ N 143°9.1’ W, Roland & D’Auria 97-055, 26 June 1997. NUTZOTIN MOUNTAINS: gravel soil on ridgetop, Copper Pass, 1943 m, 62°17.16’ N 142°31.44’ W, J. Bolivar 84-71, 28 June 1984; occasional in sterile limestone scree, Baultoff Creek, 1707 m, 62°9.13' N 141°14.51' W, C. Roland 94-144A, 27 June 1994; scattered in exposed NW-facing area of ridge, Baultoff Creek, 1707 m, 62°9.13’ N 141°14.51’ W, C. Roland 94-161, 28 June 1994; scattered on lime- stone ridge in bare mineral soil, Copper Pass, 1942 m, 62°17.16' N 142°31.44’ W, M. Cook 94344, 24 July 1994; scattered on ridge in Rhacomitrium, Klein Creek, 1747 m, 62°2.29' N 141°19.88’ W, M. Cook 95018, 15 June 1995; scattered in volcanic rubble, plateau between Bryan and Willow Creeks, 1829 m, 61°58.97' N 141°50.03’ W, M. Cook 95086, 21 June 1995; occasional on steep N-facing slope in fine scree, ridge at headwaters of Alder Creek, 1554 m, 62°28.44' N 142°15.06’ W, K.A. Beck 95-215, 30 June 1995. St. ELIAS MOUNTAINS: scattered in steep limestone gravel, butte between Dan and Copper Creeks, 1554 m, 61°21.56’ N 142°26.43’ W, C. Roland 94-230B, 14 July 1994; scattered on S-facing rock slope Mt. Chitina, 2073 m, 60°57.74' N 141°17.33' W, M. Cook 95231, 14 July 1995. WRANGELL MOUNTAINS: N-facing limestone talus slope, Lime Creek, 1646 m, 61°46.25’ N 141°50.36’ W, M. Cook 94118, 21 June 1994; scattered in frost shattered rocks on limestone ridge, Copper Pass, 1942 m, 62°17.16’ N 142°31.44' W, C. Roland 94- 283, 24 July 1994; occasional in moist organic soil betwen basalt tuff fragments, Cone Ridge, 2073 m, 62°8.21’ N 143°18.47' W, M. Cook 94371, 25 July 1994; common in stony areas, Cone Ridge, 2073 m, 62°8.21' N 143°18.47' W, C. Roland 94-311, 26 July 1994; scattered in moss on stony slopes, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24’ W, Moran & Roland 95-27, 30 June 1995; rare in moss-lichen dwarf wil- low tundra, Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85' W, M. Cook 95273, C. Roland 95-245A, 18 July 1995; scattered in barren calcareous gravel, Iron Mountain, 1868 m, 61°37.85' N 144°1.01' W, C. Roland 96-568, 16 July 1996; occasional in gravel and rubble on SE-facing limestone slopes, southwest of Alice Peak, 1628 m, 61°39.82’ N 144°7.76' W, C. Roland 96-387, 5 July 1996. This mustard was known from two localities in the Wrangell Mountains prior to our inventory: Guerin Glacier (61° 36.82’ 141° 4.98’, D. F. Murray 2103, 4 August 1968 (ALA) (Murray 1971)) and Chitistone Pass (61°37' N 141°58’ W, D. F. Murray 865, 1218, 1343, 1541, 1817 (CAN)). The collections cited above from throughout the mountain ranges in the Park connect its distribution in the Alaska Range 107 km to the northwest (Mt. Hayes Quad: Rainbow Mountain, 63°20.0' N 145°35.0’ W, G. Smith 2604, 7 July 1955 (ALA)) with stations 40 km to the east in the St. Elias Mountains, Yukon Territory (Cody 1996). This North America cordilleran species is rare in Alaska (GS $3) and in the Yukon Territory (Douglas et al. 1981). Map 129. Draba stenoloba Ledeb., Alaska Whitlowgrass — CHUGACH MOUNTAINS: mesic forb tundra, upper Tebay Lake, 1219 m, 61°12.18’ N 144°23.8’ W, M. Duffy 91114, 12 August 1991; occasional in mesic meadow and heath tundra on SW-facing slope, plateau NW of Nerelna Creek, 1372 m, 61°26.66' N 144°17.69' W, Roland & D’Auria 96-448B, 10 July 1996; occasional in well-drained forb-graminoid meadow on S-facing hill, pass between Amy Creek and Klu River, 1164 m, 61°4.01’ N 143°34.83’ W, C. Roland 96-526, 13 July 1996; Luetkea meadow underlain by glacial outwash, Falls Creek, 1006 m, 61°16.84’ N 144°29.86' W, C. Roland 96-554, 15 July 1996; steep SW-facing scree slope, Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96- 221C, 28 July 1996; few in bare soil of E-facing cut bank, 12-mile Creek, Granite Range, 1326 m, 60°50.21’ N 142°30.85’ W, C. Roland 96-785, 96- 817, 29 July 1996; occasional in moss and organic soil of herbaceous vegetation, plateau NW of Nerelna Creek, 1372 m, 61°26.66’ N 144°17.69' W, M. Cook 96303, 7 July 1996; scattered in bare min- eral soil and moss along sparsely vegetated ridge, ridge between Canyon, Pass and Tenas Creeks, 1682 m, 61°24.28' N 144°21.61' W, Cook & Losso 96323, 7 July 1996; scattered in forb herbaceous vegetation along stream, plateau between Tebay River, Hanagita River and Grant Creek, 1250 m, 61°17.65’ N 143°56.34' W, M. Cook 96368, 96411, 7 July 1996; scattered on alpine forb herbaceous slope, vic. Hanagita Peak, 1186 m, 61°4.91' N 143°38.86' W, M. Cook 96417, 96429, 7 July 1996; SE-facing rock outcrops, Upper Golconda Creek, 1311 m, 61°2.8' N 143°25.64’ W, L.A. & E.G. Viereck 11071, 7 July 1996. GULF OF ALASKA: ericaceous dwarf scrub alpine tundra, nunatak 3.5 km NW of Hanna Lake, 610 m, 60°9.95’ N 143°6.8’ W, M. Duffy 9263, 13 June 1992. NUTZOTIN MOUNTAINS: rare, with seral herbs on dry, S-facing bluff above Beaver Creek, 1097 m, 62°1' N 141911’ W, Parker & Gracz 6832, 10 August 1996; WRANGELL MOUNTAINS: occasional in protected gully draining blocky talus slope, Kuskulana Pass, 1545 m, 61°33.72' N 143°39.7' W, C. Roland 96-716, 26 July 1996. This mustard, a North American cordilleran species, was known from one locality in the Wrangell Mountains (Bonanza Ridge, 61°30’ N 142°51' W, Nordell & Schmitt 62, 238, 483, 2 July 1976 (LD & ALA) (Nordell and Schmitt 1978)) and from Yakutat Bay (F. Funston 96, 1892 (US) (Coville and Funston 1896; Hultén 1941)). The speci- mens cited above extend its range up to 230 km from the previous collections. The closest stations to our collections are 43 km to the east (Cordova Quad 60°58.0' N 145°00.0' W, C.L. Parker 1778, 12 August 1986 (ALA)) and 207 km to the north in the Alaska Range (Gulkana Quad: 62°57.0' N 145°31.0' W, D. Mesiar DCM-48, 3 July 1979 (ALA)). Map 130. Draba stenopetala Trauty., Anadyr Whitlowgrass — CHUGACH MOUNTAINS: occasional on talus slope and a ———— ee ee — > + ee ee 262 Dryas tundra, Hundell Creek, 1585 m, 61°36.75’ N 144°42.2’ W, M. Cook 93466, 28 August 1993; rocky tundra knoll on ridge crest, vic. East Fork Kiagna River, Granite Range, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-278, 29 July 1996; occasional in gravel patches, Nerelna Creek plateau, 1372 m, 61°26.66’ N 144°17.69' W, M. Cook 96307, 8 July 1996. MENTASTA MOUNTAINS: scattered in red gravel scree, SE-facing slope above Totschunda Creek, 1494 m, 62°28.28' N 142°40.5' W, Cook & Shea 96286, 3 July 1996; common in gravel on orange-colored rubble slope, Caribou Creek, 1158 m, 62°35.7' N 143°27.87' W, Roland & D’Auria 97- 017, 26 June 1997. NUTZOTIN MOUNTAINS: NW-fac- ing steep, exposed talus slope, Gold Hill, 1585 m, 62°6.73' N 141°51.82'-W, M. Cook 9270, 27 June 1992; Dryas tundra ridge between Fogenbera Pass & Shotgold Creek, 1783 m, 61°53.28’ N 141°37.1' W, M. Cook 92457, 8 July 1992; occasional on S-facing slope in Dryas-Rhacomitrium tundra, volcanic plateau between Bryan and Willow Creeks, 1829 m, 61°58.97' N 141°50.03’ W, M. Cook 95058, 18 June 1995; alpine fellfield, on ridgetop east of Mt. Drum, 1768 m, 62°4.5’ N 144°22.8’ W, K.A. Teare 1683, 4 July 1984. WRANGELL MOUNTAINS: alpine tundra, slope above Nabesna Glacier moraine, 1768 m, 61°56.6' N 143° W, M. Duffy 92113, M. Cook 9262, 26 June 1992; Dryas-sedge tundra, NW slope Mt. Sanford, 1295 m, 62°21.17' N 144°24.52’ W, M. Cook 93109, 9 June 1993; rare on E-facing gravel slope, northeast slope of Mt. Drum, 1433 m, 62°8.83' N 144°30.18' W, Cook & Roland 94054A, 11 June 1994; scattered on stony alluvium of cinder cone, N slope Mt. Sanford, 1722 m, 62°23.4’ N 144°15.9' W, C. Roland 94-004, 6 June 1994; scat- tered on steep S-facing rubble, W slope Mt. Sanford, 1615 m, 62°13.59’ N 144°26.06’ W, C. Roland 94- 042, 9 June 1994; rare in gravel of dry Salix-lichen tundra, Cooper Pass, 1942 m, 62°17.16’ N 142°31.44' W, C. Roland 94-290, 24 July 1994; scat- tered clumps on basalt tuff, Cone Ridge, 2073 m, 62°8.21' N 143°18.47’ W, M. Cook 94393, 26 July 1994; rare in bare mineral soil, Chetaslina plateau, 1615 m, 61°56.51’ N 144°25.93’ W, M. Cook 94467A, 15 August 1994; occasional in bare mineral soil of volcanic avalanche deposits, Nadina Glacier, 1768 m, 62°2.85' N 144°41’ W, M. Cook 94413, 30 August 1994; rare in gravel on moderate S-facing slope in sandy sites, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24' W, C. Roland 95-118, 1 July 1995; patchy on unstable gravel and cobble scree slope, Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85’ W, M. Cook 95274, C. Roland 95-234, 18 July 1995; occasional in gravel frost scars, Black Mountain, 1481 m, 62°20.85’ N 143°44.9' W, Cook & Beck 95147, 7 July 1995; scattered in moist, barren lime- stone gravel on gentle, NW-facing slope, 1868 m, 61°37.85' N 144°1.01' W, C. Roland 96-585A, THE CANADIAN FIELD-NATURALIST Vol. 116 - 16 July 1996; occasional in limestone gravel, ridge between headwaters of Pass and East Fork of Copper Creeks, 1792 m, 61°39.82" N’ 144°O 18 Roland 96-591, 16 July 1996; occasional in morainal deposits, valley between Ruddy Mountain and Mt. Drum, 1615 m, 62°4.61' N 144°46.39' W, C. Roland 96-325, 2 July 1996; few in limestone gravel talus, Kuskulana Pass, 1545 m, 61°33.72’ N 143°39.7' W, M. Cook 96550, 26 July 1996; rare in loose gravel on barren SW-facing slope of Snyder Peak, 1524 m, 62°4.47' N 144°30.51' W, C. Roland 96-382, 5 July 1996. This amphiberingean arctic-alpine species, unknown previously from the Park, is now known from 25 localities in the Western and Northern Wrangells, Mentasta and Nutzotin Mountains. Our collections connect its distribu- tion in the Alaska Range 147 km to the northwest (Hultén 1968) with the distribution in the Yukon Territory 105 km to the east (Cody 1996). The number of localities in the Western Wrangell Mountains may indicate that there was a local refugium from which these populations are derived. Hamilton and Thorson (1983) have suggested that the mountain slopes adjacent to Lake Ahtna in the Copper River basin could have been ice free at the end of the Wisconsin glaciation. Likewise, the populations in the Mentasta and Nutzotin Mountains, being close to the ice- free corridor, may represent areas that were ice-free earlier. This mustard is considered rare in Alaska (G3 $3S4) and in the Yukon Territory (Douglas et al. 1981). Map 131. Erysimum pallasii (Pursh) Fern., Pallas’ Wallflower — MENTASTA MOUNTAINS: bare mineral soil on S- facing limestone scree slope, Soda Lake, 1173 m, 62°32.36’ N 142°53.98' W, M. Cook 94322, 22 July 1994; occasional in steep, loose, bare rubble on S- facing slope, Devils’s Mountain, 1530 m, 62°25.62’ N 142°53.93’ W, C. Roland 96-348, 3 July 1996; few in unstable reddish gravel scree, Devils’s Mountain, 1530 m, 62°25.62’ N 142°53.93’ W, M. Cook 96279, 7 July 1996. NUTZOTIN MOUNTAINS: scattered on SW-facing rubble slope, Carl Creek, 1920 m, 62°3.52' N 141°36.27’ W, C. Roland 94- 084, 20 June 1994; rare on very unstable S-facing gravel size scree slope, Horsfeld Creek, 1768 m, 62°2.88' N 141°13.18' W, M. Cook 94150, 24 June 1994; scattered within orange hypabyssal rhyolite unit on ridge, volcanic plateau between Bryan and Willow Creeks, 1829 m, 61°58.97’ N 141°50.03’ W, M. Cook 95049, 18 June 1995; few in dry basalt gravel and sand, Wiki Creek, 1411 m, 61°54.46 N 141°10.68’ W, M. Cook 96089, C. Roland 96-066, 10 June 1996; scattered in scree just below ridge, pass between Cabin and Wiki Creeks, 1615 m, 61°53.7' N 141°11.64' W, M. Cook 3173, 14 June 1998 (ALA); scattered in unstable scree and amongst rock outcrops, Wiki Basin, 1524 m, 61°54.77' N 141°11.05’ W, M. Cook 3188, 15 July 1998 (ALA). WRANGELL MOUNTAINS: rare unstable, limestone scree, Lime Creek, 1646 m, 61°46.25’ N 141°50.36' W, M. Cook 94115, 21 June 1994. 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 131. Draba stenopetala 132. Erysimum pallasii 263 264 This circumpolar arctic-alpine species was known from two localities in the Wrangell Mountains: Russell Glacier (61°42.05’ 141°44.75’ D. F. Murray 2156, 10 August 1968 (ALA)) and Skolai River (61°37’ N 142°3' W, R. Scott s.n., 1976 (MICH) (Scott 1968)). The collections cited above extend its range 112 km to the east into the Mentasta Mountains and connect the distribution in the Alaska Range 286 km to the northwest (Healy Quad: Polychrome Pass, 63°31.0’ N 149°56.0' W, A. & R.A. Nelson 3818, 14 July 1939 (ALA)) with the range 57 km to the east in the Yukon Territory (Cody 1996). This species is considered rare in Alaska (G4 $3). Map 132. Eutrema edwardsii R. Br., Edwards’ Mock Wallflower — NUTZOTIN MOUNTAINS: Dryas tundra, ridge between Fogenbera Pass and Shotgold Creek, 1783 m, 61°53.28’ N 141°37.1' W, M. Cook 92448, 8 July 1992; scattered in wet moss of graminoid swale, volcanic plateau between Bryan and Willow Creeks, 1829 m, 61°58.97’ N 141°50.03’ W, M. Cook 95042, 17 June 1995; scattered in mesic graminoid herbaceous/shrub birch-willow scrub, Wiki Creek, 1411 m, 61°54.46’ N 141°10.68’ W, M. Cook 96065, 9 June 1996. ST. ELIAS MOUNTAINS: rare in late snow melt area on N-facing slope, wet silt with sparse forbs, Mt. Chitina, 2073 m, 60°57.74' N 141°17.33’ W, M. Cook 95250, 16 July 1995. WRANGELL MOUNTAINS: in standing water, iso- lated ridge SW of Long Glacier, 1372 m, 61°45.83’ N 144°12.12'’ W, K.A. Teare 84-86, 18 July 1984; alpine tundra, 152 m above and E of Nabesna Glacier moraine, 1768 m, 61°56.6’ N 143° W, M. Cook 92556, 26 June 1992; Dryas-sedge tundra, NW slope of Mt. Sanford, between Sanford and Boulder Creeks, 395 m, 62°21.17’ N 144°24.52’ W, M. Cook 93124, 9 June 1993; seep area in talus on rock out- crop, Jaegar Mesa, 1893 m, 62°15.9' N 143°1.24’ W, Moran & Roland 95-31, 30 June 1995; patchy in moss next to rivulet with dwarf willows and alpine forbs, Crystalline Hills, 1585 m, 61°23.59' N 143°31.85' W, M. Cook 95279, 18 July 1995; poly- gon tundra, Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5' W, Potkin & Leggett 95-069C, 29 July 1995; moist meadow intermixed with clumps of Salix lanata, Nikolai Pass, 1280 m, 61°26’ N 142°40' W, Batten & Barker 96-183B, 26 July 1996 (ALA). This circumpolar arctic-alpine species was previously known from two localities in the Wrangell Mountains: Russell Glacier (61° 41.88’ N 141° 44.49’ W, D. F. Murray 2203, 11 August 1968 (ALA)) and Chitistone Pass/upper Skolai River Valley, 61°37’ N 142°3’ W, R. Scott s.n., 1967 (MICH) (Scott 1968) and D. F. Murray 1035, 21 July 1967 (CAN)). The new collections cited above extend its range 148 km west into the Wrangells and 87 km south into the St. Elias Mountains and connect the distribution in the Alaska Range 180 km to the northwest (Healy Quad: Cantwell, 63°24.0’ N 148°56.0' W, L.J. Palmer 1931, 10 July 1927 (ALA)) with the distribution 137 km to the east in the Yukon Territory (Cody 1996). Map 133. THE CANADIAN FIELD-NATURALIST Vol. 116 Halimolobos mollis (Hook.) Rollins, Soft Fissure- wort — CHUGACH MOUNTAINS: Third Lake, 4.8 km N of Chitina, 396 m, 61°34.12’ N 144°26.37’ W, M. Cook 8962, 4 June 1989. MENTASTA MOUNTAINS: occasional in disturbed sites on dry, S-facing slopes in sagebrush-grass vegetation, SW end of Devil's Mt., 942 m, 62°24.95'’ N 142°54.86’ W, C. Roland 96-233, M. Cook 96240, 22 June 1996. WRANGELL MOUNTAINS: common in dry, rocky areas on steep SE-facing slope in sagebrush-grass vegetation, Nabesna River, 0.6 km S of Bond Creek, 1106 m, 62°15.05’ N 142°54.68' W, C. Roland 96-165, M. Cook 96188, 19 June 1996. This North America arctic-alpine mustard was collected in the vicinity of the Park at Copper Center (W.L. Poto 14, 1 June 1902 (US) (Poto 1902*; Hultén 1941-1950); C.W. Heidemann 31, 1908 (US) (Hultén 1941-1950)) and at Chitina (J.P. Anderson 2017, 1935 (S) (Hultén 1941- 1950)). The new collections cited above along the Nabesna River are 130 km north of the Copper Center station and 112 km south of a station near the Robertson River (O.J. & M.E. Murie, 1921 (US) (Hultén 1941-1950)). Map 134. Lesquerella arctica (Wormsk.) S. Wats., Arctic Bladderpod — CHUGACH MOUNTAINS: SE-facing bluff above road and W of bridge, confluence of Chitina and Copper Rivers, 274 m, 61°31.55’ N 144°24.85’ W, M. Cook 93102, 21 May 1993; SW-facing scree, Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-297, 31 July 1996. MENTASTA MOUNTAINS: few localized on S-facing limestone talus slope sparsely vegetated with midgrass-herbs, Soda Lake, 1173 m, 62°32.36' N 142°53.98’ W, M. Cook 94323, 22 July 1994; river terrace, Totschunda Creek, 725 m, 62°26.94' N 142°40.8' W, C. Roland 96-250, 23 June 1996. St. ELIAS MOUNTAINS: dry benchland meadow, Chitina River, 335 m, 61°9.02’ N 142°39.95’ W, D. Miquelle 84-10, 20 June 1984; few localized at edge of active flood zone on sand spit 0.8 km N of Jakes Bar, Chitina River, 93 m, M. Cook 94230, 9 July 1994 (ALA); few localized in dry sandy clay mineral soil on ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94' N 141°6.75' W, M. Cook 95221, 13 July 1995. WRANGELL MOUNTAINS: scattered on limestone outcrop above mixed Picea glauca-Poplar forest, Crystalline Hills, 618 m, 61°23.1’ N 143°36.01’ W, M. Cook 3146, 24 June 1998. This circumpolar arctic-alpine bladderpod was collected at Chitina (61°31.55’N 144° 27.89’W, L. Spetzman 2557 (CAN) (Rollins 1973); D.F. Murray 6161, 6232 (ALA)) and on the upper Chitina River (61°01.84N’ 141°37.27'W, HM. Laing 91, 92 (CAN) (Porsild 1939, Hultén 1941-1950)). When A.E. Porsild classified Laing’s collection, he deter- mined these specimens to be L. arctica var. purshii Wats. (Porsild 1939). However, Hultén (1945), Rollins (1973) and Cody (1996) do not recognize this variety. This plant has a restricted distribution in the Park but with a wide ecological tolerance. It is now known from eight localities along the Chitina River and in the Mentasta Mountains on flood- plains, alpine scree and dry south-facing bluffs. The collec- tions cited above extend its range 183 km to the north into 2002 CooK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 136. Smelowskia borealis 265 - -+-ooOo oe dC +--+ —-—-—-- —- 266 the Mentasta Mountains. These collections also connect the distribution between the Alaska Range 243 km to the north- west (Rapids, 63°37.56'N 145°53.06'W, O.J. Murie s.n., 7 June 1992 (US), E. Scamman 216 (G, S) (Hultén 1941- 1950)) with the distribution 188 km to the east in the Yukon Territory (Cody 1996). Map 135. Smelowskia borealis (E. L. Greene) Drury & Rollins, Northern Smelowskia — MENTASTA MOUNTAINS: few on sparsely vegetated scree slope, Totschunda Creek, 1494 m, 62°28.28’ N 142°40.5’ W, M. Cook 96296, 7 July 1996; few in unstable limestone scree, Lost Creek, 1646 m, 62°34.58’ N 143°5.58’ W, C. Roland 94-265, 23 July 1994; locally abundant in limestone gravel, Trail Creek, 1615 m, 62°36.05’ N 143°17.63' W, C. Roland 95-012, 6 June 1995. NUTZOTIN MOUNTAINS: rare on S-facing granidorite boulder-cobble slope, Carl Creek, 1859 m, 62°3.07' N 141°35.01’ W, M. Cook 94129, 22 June 1994; scattered on steep, unstable S facing fine scree slope, Horsfeld Creek, 1768 m, 62°2.88’ N 141°13.18’ W, M. Cook 94159, 24 June 1994. WRANGELL MountTAINs: N-facing scree slope, Lime Peak, 1707 m, 61°28.41' N 141°29.44’ W, M. Duffy 92144, 7 July 1992; occasional in loose limestone and green- stone rubble, Alice Peak, 1628 m, 61°39.82' N 144°7.76' W, C. Roland 96-396A, 5 July 1996; occa- sional in gravel 15 m below ridge, Elliot Creek, 1926 m, 61°38.99’ N 144°3.51' W, M. Cook 96470, 7 July 1996; steep unstable scree of loose shaley orange-tan rocks, Nikolai Pass, 1280 m, 61°26’ N 142°40’ W, Batten & Barker 96-024, 23 July 1996. This species is an Alaska-Yukon endemic with an arc- tic-alpine distribution. Hultén (1942) indicated that it was a remnant of a tertiary type that survived only in unglaciated areas. It was known from four localities in the Wrangell-St. Elias Mountains: Russell Glacier (61°42.0’ N 141°45.0’ W, D. F. Murray 2180, 11 August 1968 (ALA)); Guerin Glacier (61°37.0' N 141°05.0’ W, D. F. Murray 2096, 4 August 1968 (ALA)); Sheep Glacier (61°42.0’ N 141°39.0’ W, D. F. Murray 2254, 15 August 1968 (ALA)) and Chitistone Pass/upper Skolai River Valley (61°37' N 142°3' W, 604 m, R. Scott 2238, 1967 (MICH) (Scott 1968)). A collection in Hultén (1968) on the upper Copper River is that of W.L. Poto from the East Fork of the Chistochina (62°53.78'N 144°40.20’W, #157, 22 August 1902 (US) (Poto 1902*)). The Chistochina River was mapped incorrectly in Hultén (1941) and we were not able to locate an annotation for the station that appears on the upper Copper River in Hultén (1968). The collections cited above extend the range of this species 124 km north into the Mentata Mountains, 111 km west into the western Wrangell Mountains and connect the distribution 88 km to the northwest in the Alaska Range (Mt. Hayes Quad: Rainbow Mountain, 63° 13’ N, 145° 38’ W, 63°15.0' N 145°35.0' W, C.L. Parker 1734, 2 September 1985 (ALA)) with the range to the east in the Yukon Territory (Cody 1996). Map 136. Smelowskia calycina (Stephan) C.A. Meyer s.1., False Candytuft — NuTZOTIN MOUNTAINS: S-facing scree slope, knob 2.4 km W of Ptarmigan Creek, 1494 m, 61°55.55' N 141°6.72' W, M. Cook 92467, THE CANADIAN FIELD-NATURALIST Vol. 116 M. Duffy 92185, 9 July 1992; few in loose rhyolite gravels at edge of Dryas/graminoid tundra stringers on W-facing slope, Wiki Creek, 1433 m, 61°53.9' N 141°9.51’ W, C. Roland 94-109, 22 June 1994; sparsely vegetated SW-facing scree slope, Horsfeld Creek, 1768 m, 62°2.88’ N 141°13.18’ W, M. Cook 94156, C. Roland 94-130, 24 June 1994 (ALA); scattered on rhyolite talus, plateau between Bryan and Willow Creeks, 1829 m, 61°58.97' N 141°50.03’ W, M. Cook 95043, 17 June 1995; very rare in grav- els on dry tundra knob, Wiki Creek, 1411 m, 61°54.46' N 141°10.68’ W, C. Roland 96-059, 96- 076, 8 June 1996; few scattered in scree, Wiki Basin, 1524 m, 61°54.77' N 141°11.05’ W, M. Cook 3185, 15 July 1998 (ALA); scattered in scree, Sonja Creek, 1494 m, 61°57.14' N 141°21.43’ W, M. Cook 3193, 16 July 1998 (ALA). WRANGELL MOUNTAINS: N-fac- ing scree slope, Lime Creek, 1707 m, 61°47.2’ N 141°49.06’ W, M. Cook 92509, 92512, M. Duffy 92145, 7 July 1992; Dryas sedge tundra, NW slope of Mt. Sanford between Sanford and Boulder Creeks, 395 m, 62°21.17’ N 144°24.52’ W, M. Cook 93121, 9 June 1993; rare in rocks on moderate S-fac- ing slope, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24’ W, C. Roland 95-117A, 1 July 1995. This species was collected in the vicinity of Russell Glacier (61°41.64'N 141°44.85'W, D.F. Murray 2200, 11 August 1968 (ALA) (Murray 1971)). The new collections cited above extend its range 156 km to the east into the western Wrangell Mountains. The closest stations to those in the Wrangell and Nutzotin Mountains are 405 km to the west in the McGrath Quad (Farewell Mountain, 62°31.0'’ N 153°43.0’ W, C. Parker 900, 11 July 1983 (ALA)), 439 km to the northwest in the Tanana Quad (Moose Creek, 65°36.0’ N 150°41.0' W, K.C. Kassler 282, 25 June 1979 (ALA)) and 323 km to the northeast in the Yukon Territory (Cody 1996). Five of the specimens cited above were determined as S. calycina (Stephan) C.A. Meyer var. porsildii Drury & Rollins based on the presence of all entire, linear leaves. However these individuals came from populations that also had plants with either all divided leaves or both types of leaves. In our area, leaf shape appears to be a plastic char- acter, whereas populations in unglaciated Alaska, eastern- most Siberia and Western North America are more consis- tent with regards to the simple leaf shape (Hultén 1944). Hultén commented that, “This species is a fine example of the geographical conditions in species older than the Pleistocene glaciation” and suggested that the simple- leaved form (S. calycina var. porsildii) survived glaciation whereas the divided leaf form is a derived state. The occur- rence of both forms in the Nutzotin and Wrangell Mountains, populations that are 800 km disjunct from the main arctic range, may indicate that these areas were ice- free earlier. Map 137. Subularia aquatica L., Awlwort — WRANGELL MOUNTAINS: common along muddy shoreline, Fox Farm Lake, 727 m, 62°19.98’ N 144°50.02’ W, M. Cook 95311, 95303, C. Roland 95-252, 25 July 1995; gravel bottom, Chelle Lake, 930 m, 62°11.56'N, 144°51.97'W, M. Cook 3606, 1 September 2000. 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 267 This circumpolar boreal-montane aquatic mustard is of restricted distribution in Alaska and rare in the Yukon Territory (Douglas et al. 1981). The stations in the Wrangell Mountains cited above are 127 km south of the closest station in the Alaska Range near Summit Lake (A.E. & R.T. Porsild 546, July 1926 (CAN) (Hultén 1941-1950)). Map 138. Thlaspi arcticum Pors., Arctic Pennycress — WRANGELL Mountains: NE-facing scree slopes, Iron Mountain, 1676 m, 61°37.64’ N 144°1.42’ W, M. Duffy 91150, 27 August 1991, C. Roland 96-590, M. Cook 96465, 16 July 1996. | This Alaska- Yukon endemic with an arctic-alpine distri- bution is one of the rarest plants in the Park, known only from the one locality cited above. It is rare in Alaska (G3 S3) and in the Yukon Territory (Douglas et al. 1981). Most localities are widely disjunct in Alaska and the Yukon and represented by very few individuals (Murray and Lipkin 1987). The collection in the southwest Wrangells cited above is 252 km northeast of a station in the Anchorage Quad (North Campbell Creek Canyon, 61°07.17' N 149°29.75' W, Lichvar et al. 8085, 19 July 1994 (ALA)), 318 km southeast of a station in the Healy Quad (Windy Creek, 63°24.0’ N 149°17.0’ W, D. Heebner 100.2, 7 July 1977 (ALA)) and 303 km west of a station in the Yukon Territory (Cody 1996). Map 139. SAXIFRAGACEAE Mitella pentandra Hook., Alpine Mitrewort — GULF OF ALASKA BASIN: tall forb herbaceous alpine tundra, Robinson Mountains, 671 m, 60°5.64'’ N 142°17.17’ W, M. Cook 9233, M. Duffy 92214, 12 June 1992. ST. ELIAS MOUNTAINS: mixed forb herbaceous mead- ow, Icy Bay, 427 m, 60°0.76’ N 141°27.1' W, K. Beck s.n., 12 August 1987; heather tundra, Floral Pass, 549 m, 59°57.91’ N 139°57.76' W, M. Cook 87-146, 21 August 1987; snowflush streamlet, Samovar Hills, 2200 m, 60°4.15’ N 140°28.75’ W, M. Cook 92522, 18 July 1992. WRANGELL MOUNTAINS: tall forb-graminoid herbaceous mead- Ow, iakinaGlacier, 1219 m,; 61°33.64'’ N 143°19.01’ W, M. Cook 96531, C. Roland 96-704, 24 July 1996. This species is North American with a cordilleran distri- bution and is rare in the Yukon Territory (Douglas et al. 1981). The collections cited above extend its range 212 km northeast into the Wrangell Mountains and 348 km east into the southern St. Elias mountains and connect the range 166 km to the east in the Yukon Territory (Cody 1996). Map 140. Saxifraga adscendens L. ssp. oregonensis (Raf.) Baciagalupi, Small Saxifrage — CHUGACH MOUNTAINS: occasional in serpentinized ultramafic unit on ridge S of Chakina River, 1768 m, 61°6.19' N 142°55.03' W, M. Cook 94229, 8 July 1994; rocky ridge crest and W-facing slide rock, West Fork Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-294, 29 July 1996 (ALA); scattered on moist gravel slope, Nelson Mountain, 1558 m, 61°19.37' N 143°48.83' W, M. Cook 96370, 7 July 1996. St. ELIAS MOUNTAINS: rare in solifluction lobes, limestone ridge between Dan & Copper Creeks, 1554 m, 61°21.56’ N 142°26.43’ W, M. Cook 94293, 14 July 1994. WRANGELL MOUNTAINS: scat- tered in dry, rocky limestone talus, Lakina Glacier, 1219 m, 61°33.64’ N 143°19.01' W, M. Cook 96537, C. Roland 96-705, 25 July 1996; few in fine gravels, in greenstone unit, Iron Mountain, 1868 m, 61°37.85’ N 144°1.01’ W, M. Cook 96467, 7 July 1996; bare organic soil and moss, Grotto Creek, 1847 m, 61°30.56’ N 142°24.79' W, M. Cook 96510, 7 Suly 1996; scattered in moss and organic soil of limestone outcrops, Lakina Glacier, 1219 m, 61°33.64’ N 143°19.01' W, M. Cook 96537, 7 July 1996. This North American cordilleran saxifrage was known from one locality in the Wrangell Mountains (Chitistone Pass, 61°37’ N 141°58’ W, 1979 m, D. F. Murray 683, 1064 (CAN)). The collections cited above extend its range 107 km into the Western Wrangells and 103 km south into the Chugach Range. These collections join the cordilleran range 240 km to the southwest in the Anchorage Quad (Nike Site, 61°15.70' N 149°32.25’ W, Duffy et al. 381, 29 June 1994 (ALA)) with the distribution 161 km to the east in the Yukon Territory (Cody 1996). Map 141. Saxifraga bracteata D. Don (S. sibirica L.), Siberian Saxifrage — NUTZOTIN MOUNTAINS: occasional on rock-Dryas-Rhacomitrium SW-facing slope, Carl Creek, 1920 m, 62°3.52' N 141°36.27' W, Cook & Roland 94091A, 20 June 1994. St. ELIAS MOUNTAINS: dry forb alpine herbaceous talus, Samovar Hills, 610 m, 60°8.35’ N 140°39.6' W, M. Cook 87-77, 24 August 1987. The specimens cited above extend the range of this amphiberingean saxifrage 259 km north into the Nutzotin Mountains and 180 km east to the Samovar Hills from Kayak Island (59°56.0’ N 144°23.0’ W, Cunningham et al. 124-78, 1 June 1978 (ALA)). These collections connect the Pacific Coastal range to the southwest with the range 348 km to the southeast (Hultén 1968). Map 142. Saxifraga foliolosa R. Br., Leafystem Saxifrage — NUTZOTIN MOUNTAINS: occasional in volcanic ash, upper Flat Creek, 1323 m, 61°58.97' N 141°41.25’ W, C. Roland 95-045, 17 June 1995. WRANGELL MOUNTAINS: sedge tundra frost boil, Cheshnina Plateau, 1640 m, 61°50.33’ N 144°22.5' W, M. Duffy 91087, 21 July 1991; scattered in moist organic soil, Mesa Creek, 2073 m, 62°8.21' N 143°18.47' W, M. Cook 94369, C. Roland 94-294, 25 July 1994; bare moist mineral soil, Chetaslin Ridge, 1615 m, 61°56.51' N 144°25.93' W, M. Cook 94483, 16 August 1994; wet meadow, Jaegar Mesa, 1893 m, 62°15.9' N 143°1.24’ W, Moran & Roland 95-46, | July 1995; Lakes Plateau, 1890 m, 62°4.4’ N 143°23.5' W, M. Potkin 95-148, 30 July 1995. This saxifrage is circumpolar with an arctic-alpine dis- tribution. The stations cited above connect the range 110 km to the southwest at Thompson Pass (Valdez Quad, 61°08.0' N 145°44.0' W, C.L. Parker 2362, 21 July 1990 (ALA)) with the range 141 km to the north in the Alaska Range in the Mt. Hayes Quad (Upper Berry Creek, 63°30.0' N 144°35.0' W, L.A. Spetzman 1038, 5 August —— ewe CS -C ” 268 THE CANADIAN FIELD-NATURALIST Vol. 116 140. Mitella pentandra COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 141. Saxifraga adscendens ssp. oregonensis 144. Chamaerhodos erecta ssp. nuttallii 269 am i nee ee tee a eee 270 1957 (ALA)). It is considered rare in the Yukon Territory by Douglas et al. (1981). Map 143. ROSACEAE Chamaerhodos erecta (L.) Bunge ssp. nuttallii (Torr. & Gray) Hult., Little Rose — MENTASTA MounrtTaAINs: locally abundant in bare soil on steep, S-facing slopes, Devil’s Mountain, 942 m, 62°24.95' N 142°54.86' W, C. Roland 96-225, M. Cook 96237, 22 June 1996; occasional on gravel scree slope, NE slopes of Devils’s Mountain, 1530 m, 62°25.62' N 142°53.93' W, M. Cook 96284, 3 July 1996. This amphiberingian species is disjunct in the Mentasta Mountains from the Anchorage Quad, 242 km to the south- west (Mile 74 Glenn Highway, 61°46.0’ N 148°33.0’ W, S.L. Welsh 5001, 30 July 1965 (ALA)), the Eagle Bluff Quad 285 km to the north’ (64°50.0' N 141°11.0’ W, R. Lipkin 80-22, 11 June 1980 (ALA)) and Kluane Lake in the Yukon Terriotry 194 km to the east. Two additional small populations of this amphiberigian species have been observed by M. Cook, one 7 km up the Nabesna River from the Devil's Mountain locality at the confluence with the Jacksina River (62°21.51’N 142°52.87'W) and one 8 km east and across the Nabesna River on a river terrace at Cabin Creek (62°23.57'N 142°46.74'’W). Map 144. Potentilla arguta Pursh ssp. convallaria (Rydb.) Keck, Tall Cinquefoil — CHUGACH MOUNTAINS: few on dry S-facing slope, Granite River, Granite Range, 884 m, 60°44.63’ N 142°6.05’ W, M. Cook 96610, 7 July 1996. The specimen cited above is an extension of the range of this North American cinquefoil, 209 km to the east of a sta- tion near Thompson Pass (Hultén 1968), 244 km southeast of a station near Slana (Hultén 1968) and 207 km west of its distrubution in the Yukon Territory (Cody 1996). Map 145. Potentilla biflora Willd., Two-Flower Cinquefoil — CHUGACH MOUNTAINS: rare in dwarf scrub tundra, Chakina River, 1768 m, 61°6.19’ N 142°55.03’ W, M. Cook 94227, 8 July 1994; locally common in Dryas-graminoid tundra, Verde Ridge, 1554 m, 61°14.03' N 143°28.52' W, Roland & D'Auria 96- 483, 11 July 1996; occasional in Dryas-lichen tun- dra, Nerelna Creek plateau, 1372 m, 61°26.66’ N 144°17.69' W, M. Cook 96306, 7 July 1996; occa- sional in Dryas-lichen tundra, Nelson Mountain, 1558 m, 61°19.37' N 143°48.83’ W, M. Cook 96375, 7 July 1996. MENTASTA MOUNTAINS: Dryas forb tun- dra, limestone ridge between Lost and Platinum Creeks, 1646 m, 62°34.58’ N 143°5.58’ W, M. Cook 94339, 23 July 1994; upper Trail Creek, 1341 m, 62°37.09’ N 143°16.06’ W, M. Potkin 95-042, 26 July 1995; scattered in Dryas-lichen-moss stringers, Lost Creek, 1722 m, 62°36.45’ N 143°12.03’ W, M. Cook 95179, 5 July 1995; locally abundant in argillite scree, Totschunda Creek, 1561 m, 62°30.62' N 142°47.21' W, C. Roland 96-365, 4 July 1996. NUTZOTIN MOUNTAINS: alpine tundra, Sheep Creek, 1554 m, 62°8.61' N 141°46.93’ W, J. Bolivar 84-61, 26 June 1984; dwarf scrub sedge tundra, Solo Mt., THE CANADIAN FIELD-NATURALIST Vol. 116 1620 m, 62°11’ N 143°31.42'’ W, M. Cook 92052, 8 July 1992; scattered in SE-facing limestone, Baultoff Creek, 1707 m, 62°9.13’ N 141°14.51’ W, Cook & Roland 94191, 28 June 1994; scattered in bare min- eral soil, Cooper Pass, 1942 m, 62°17iG@ on 142°31.44’ W, M. Cook 94342, 24 July 1994. Sr. ELIAS MOUNTAINS: common on N & S facing slopes in Dryas tundra & on ledges and crevices of lime- stone, ridge between Dan & Copper Creeks, 1554 m, 61°21.56’ N 142°26.43' W, M. Cook 94285, 13 July 1994; occasional in Dryas-sedge-lichen tundra, Mt. Chitina, 2073 m, 60°57.74’ N 141°17.33’ W, M. Cook 95256, 16 July 1995. WRANGELL MOUNTAINS: moist silt and humus, ridge SW of Long Glacier, 1372 m, 61°45.83' N 144°12.12' W, K.A. Teare 84- 77B, 18 July 1984; N-facing scree slope, Lime Peak, 1707 m, 61°28.41' N 141°29.44' W, M. Duffy 92208, 11 July 1992; Dryas-sedge tundra, Iron Mountain, 1676 m, 61°22.15' N 144°1.4’ W, M. Cook 92516, 11 July 1992; Dryas-sedge tundra, NW slope of Mt. Sanford between Sanford and Boulder Creeks, 395 m, 62°21.17' N 144°24.52’ W, M. Cook 93117, 9 June 1993; W-facing slope, volcanic ridge between upper Jacksina River and Mesa Creek, 2073 m, 62°8.21' N 143°18.47' W, C. Roland 94-318A, 27 July 1994; tundra, Jaegar Mesa , 1893 m, 62°15.9' N 143°1.24’ W, Moran & Roland 95-57, 3 July 1995; dry tundra near ridge crest, Nikolai Pass, 1280 m, 61°26’ N 142°40’ W, Batten & Barker 96-046, 23 July 1996; occasional in gravel on limestone ridge, Alice Peak, 1628 m, 61°39.82’ N 144°7.76’ W, C. Roland 96-389, 5 July 1996; few at edge of Dryas- graminoid heath adjacent to limestone cliff, headwa- ters of Grotto Creek, 1847 m, 61°30.56’ N 142°24.79' W, M. Cook 96511, 7 July 1996; scat- tered in limestone gravel talus, Kuskulana Pass, 1545 m, 61°33.72' N 143°39.7’ W, M. Cook 96551, 7 July 1996. This amphiberingian arctic-alpine species was previously known from only two localities in the Wrangell Mountains: between the Sanford River and Slana on the Copper River trail (62°24.70 144°5.68, 1372 m, W.L. Poto s.n., 30 July 1902 (US) (Poto 1902*; Hultén 1941—1950)) and Frederika Glacier (61°41.95 142°12.59, 1128 m, R. Scott 2651, 7 July 1968 (ALA) (Scott 1968)). It is now known throughout the mountains of the Park. The specimens cited above extend its range 114 km south into the Chugach Mountains and 72 km north into the Nutzotin Mountains. Map 146. Potentilla diversifolia Lehm., Varileaf Cinquefoil — CHUGACH MOUNTAINS: mixed forb meadow, Flood Creek, 61°42.88’ N 141°58.65’ W, M. Duffy 91061, 16 July 1991; subalpine meadow, Towhead Mountain, 1433 m, 61°3.21’ N 142°39.8' W, Duffy & Barnes 96-164, 96-174, 7 August 1996; willow thickets, Granite River, 823 m, 60°44’ N 142°13’ W, Batten & Barker 96-362, 30 July 1996; occasional in mixed herbaceous meadow, Nerelna Creek Plateau, 1372 m, 61°26.66’ N 144°17.69' W, M. Cook 96301, 96409, 7 July 1996; forb herbaceous CooK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 148. Potentilla drummondii 271 22 meadow, Lake Creek, 975 m, 61°11.4' N 143°49.45’ W, M. Cook 96403, 7 July 1996; mixed forb meadow, Hanagita Peak, 1186 m, 61°4.91' N 143°38.86' W, M. Cook 96420, 7 July 1996; forb meadow, W. Fork Goat Creek, 1487 m, 60°59.8’ N 142°11.8' W, M. Cook 96585, 28 July 1996; Upper Golconda Creek, 1311 my 61°2.8' N 143°25.64 6 WOE vA, ee EG. Viereck 11069, 7 July 1996; occasional in ericaceous dwarf scrub tundra, Juniper Island, 1346 m, 60°36.17' N 142°14.96' W, M. Cook 94233, 10 July 1994; Goat Creek, 1487 m, 60°59.5’ N 142°1' W, Batten & Barker 96-273, 29 July 1996. St. ELIAS MOUNTAINS: meadows and tundra, Lime Butte, 1554 m, 61°21.56’ N 142°26.43' W, C. Roland 94-238A, 13 July 1994. WRANGELL MOUNTAINS: mesic graminoid-forb mead- ow, Chetaslina Ridge, 1615 m, 61°56.51’ N 144°25.93' W, M. Cook 94486, C. Roland 94-344A, 17 August 1994; ridge SW of Long Glacier, 1372 m, 61°45.83’ N 144°12.12’ W, C. Teare 84-76B, 18 July 1984; steep E-facing meadow, Nikolai Mine, ‘1695 m, 61°27’ N 142°39’' W, Batten & Barker 96-077, 24 July 1996; graminoid meadow, Boulder Lake, 1036 m, 62°31.27' N 144°11.26' W, M. Cook 95361, 2 August 1995. This species is North American with a cordilieran distribution. The collections cited above connect the range 70 km to the west on the Richardson Highway (Hultén 1968) with the range 141 km to the south- east in the Yukon Territory. Map 147. Potentilla drummondii Lehm., Drummond’s Cinquefoil — CHUGACH MOUNTAINS: mesic forb tun- dra, Upper Tebay Lake; 1219 im, 61°92.18" N 144°23.8' W, M. Duffy 91109, 12 August 1991; occasional in dry graminoid meadow, Granite River, Granite Range, 884 m, 60°44.63’ N 142°6.05’ W, M. Cook 96607, 7 July 1996; scattered in ericaceous- graminoid meadow, vic. 12-mile Creek, Granite Range, 1271 m, 60°48.85’ N 142°33.52’ W, M. Cook 96632, 7 July 1996; scattered between willow, Upper Falls Creek, 655 m, 61°14.17' N 144°28.24’ W, L.A. & E.G. Viereck 11038, 7 July 1996; mixed mesic forb herbaceous snow beds, Middle Fork of the Bremner River, 869 m, 60°55.05’ N 143°43.86' W, Duffy & Barnes 96-083, 6 August 1996; few at margin of bluejoint herb meadow, Upper Tebay Lake, 579 m, 61°11’ N 144°24' W, Parker & Gracz 6739, 8 August 1996. St. ELIAS MOUNTAINS: moist herbaceous patches among birch shrubs, upper Chitina River, 503 m, 61°5.35’ N 141°56.3’ W, Parker & Duffy 6704, 6 August 1996. This North American cordilleran species is disjunct in Alaska from southern British Columbia, southwest Alberta, the Cascade Mountains of Oregon, Washington and Northern California (Hitchcock and Cronquist 1961). There are only two other known localities of this species in Alaska which are 240 km to the west of the specimens cited above from the Chugach Mountains (Seward Quad: Turnagain Pass, 60°48.0' N 149°11.0' W, J. Ver Hoef 135, 18 July 1979 (ALA); Anchorage Quad: Hatcher Pass, 61°47.0' N THE CANADIAN FIELD-NATURALIST Vol. 116 149°17.0' W, C.L. Williams 89110, 4 August 1989 (ALA)). It is considered rare in Alaska (G1 S1). Map 148. Potentilla litoralis Rydb. (P. virgulata Nels., P. pennsylvanica var. virgulata (A. Nels.) T. Wolf), Pennsylvania Cinquefoil — CHUGACH MOUNTAINS: scattered in Dryas dwarf shrub tundra on open gravel bars, Tana River floodplain, 335 m, 61°11’ N 142°51' W, Parker & Gracz 6811, 9 August 1996. MENTASTA MOUNTAINS: dry rocky soil on steep S slope, between Platinum and Totschunda Creek, 1280 m, 62°27.83'’ N 142°46.23' W, A.R. Batten 98- 13. NUTZOTIN MOUNTAINS: scattered in open low shrub birch, disturbed sandy soil of airstrip, Horsfeld airstrip, 1097 m, 62°1’ N 141°11’ W, Parker & Gracz 6893, 12 August 1996. ST. ELIAS MOUNTAINS: occasional in S-facing gravels and open ground with seral herbs, ridge between Logan and Walsh Glaciers, 1951 m, 60°53.94’ N 141°6.75’ W, C. Roland 95-154, 11 July 1995; occasional on dry, dis- turbed SE-facing gravel slopes, Mt. Chitina, 2073 m, 60°57.74’ N 141°17.33’ W, C. Roland 95-212, 15 July 1995. This North American arctic-alpine species was collected on the upper Chitina River (61°1.32 N 141°37.69 W; H.M. Laing 113, 1925 (CAN) (Porsild 1939, Hultén 1941—1950)) and near Slana (62°36.10’N 143°45.97W; Viereck & Richter 10378, 26 July 1980 (ALA)). The specimens cited above extend its range 150 km east into the Nutzotin Mountains and 70 km west down the Chitina River. These new collections connect the distribution between the Kenai Peninsula 404 km to the west (Hultén 1968), the Alaska Range 288 km to the north (Hultén 1968) and the Yukon Territory 41 km to the east (Cody 1996). Map 149. Potentilla rubricaulis Lehmann (P. hookeriana Lehm. ssp. hookeriana var. furcata (Pors.) Hult.), Rocky Mountain Cinquefoil — CHUGACH MOUNTAINS: rare in Dryas-graminoid tundra, between Granite and Lime Creeks, 1829 m, 61°0.22’ N 141°51.1’ W, C. Roland 96-751, 28 July 1996. This species is North American with an arctic-alpine distribution. The locality cited above is 900 km south of its range in northern Alaska (Philip Smith Mountain Quad: Mt. Hultén, 68°27.0' N 149°18.0’ W, D.F. Murray 9019, 2 August 1977 (ALA)) and 235 km east of collections in the Yukon Territory (Cody 1996). Map 150. FABACEAE Astragalus adsurgens Pall. ssp. viciifolius (Hult.) Welsh (A. adsurgens var. tananaicus (Hultén) Barneby) — MENTASTA MOUNTAINS: semi-prostrate mats on gravel bar, Nabesna River at Totschunda Creek, 725 m, 62°26.94' N 42°40.8' W, C. Roland 96-251, M. Cook 96252, 23 June 1996 (ALA). NUTZOTIN MOUNTAINS: mixed forb herbaceous vege- ‘tation in gravel, Chisana airstrip, 1006 m, 62°4.16' N 142°2.84’ W, M. Duffy 92190, 9 July 1992. This Alaska-Yukon endemic with a boreal-montane dis- tribution had been collected near Chitina (Kotsina River floodplain, 61°32.23’ N 144°23.63' W, J. Major s.n., 31 July 1980 (ALA); Chitina River Campground, 61°31.06’N 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 151. Astragalus adsurgens ssp. viciifolius 152. Astragalus eucosmus ssp. sealei 273 274 144°23.67' W, Khokhryakov et al. 6102, 8 July 1981 (ALA)). The new collections cited above are 140 km north of the Chitina collections and extend the range of this species into the Mentasta and Nutzotin Mountains. The closest collections to those cited above are 85 km to the north at mile 1247 on the Alaska Highway (J.P. Anderson 9188 (S) (Hultén 1946)) and 137 km to the east in the Yukon Territory. This subspecies is known only from Alaska and the Yukon as indicated on the distribution map, whereas A. adsurgens sensu lat is widely distributed in the western plains and mountains of the United States, Canada and in Asia. Map 151. Astragalus eucosmus Robins. ssp. sealei (Lepage) Hultén, Elegant Milk-Vetch — MENTASTA MOUNTAINS: scattered in stony tundra on limestone, ridge between Little Jack and Trail Creeks, 1615 m, 62°36.05’ N 143°17.63" W,.C. Reland 95-019, 7 June 1995. St. ELIAS MOUNTAINS: scattered on SE- facing limestone rubble slope, Mt. Natazhat, 1716 m, 61°35.38’ N 141°1.83' W, C. Roland 95-062, 19 June 1995. There were two stations of this Alaska- Yukon endemic mapped by Hultén (1968) in the Park. The station at Slana may have been Hultén’s collection since he collected in this vicinity in 1960. We have not been able to find data for the station in the central Wrangells which could have been W. L. Poto’s from the 1902 U.S.G.S. expedition (Mendenhall and Schrader 1903; Poto 1902*). However, Hultén includ- ed the rest of Poto’s collections in his 1941-1950 flora so it seems unlikely that he would have missed this specimen. The specimen from Mt. Natazhat cited above is 195 km east of the previous collections and extends the range of this species into the St. Elias Mountains. Our collections are 213 km southeast of a station in the Alaska Range (Healy Quad: 63°24.0' N 148°56.0’ W, G. Smith 2201, 4 September 1953 (ALA)) and 178 km west of a station in the Yukon Territory (Cody 1996). Astragalus eucosmus sensu lat ranges from Labrador, Newfoundland, Alaska and the Yukon Territory, south to Maine and west to Utah and Colorado. Map 152. Astragalus harringtonii (Rydb.) Hultén (A. robbinsii (Oakes) Gray var. harringtonii (Rydb.) Barneby, A. robbinsii (Oakes) Gray ssp. harringtonii (Rydb.) Hult.), Harrington Milk-vetch — CHUGACH MOUNTAINS: occasional near edge of forest in erod- ing soils, Tana Dunes, 488 m, 61°6.19’ N 142°55.03’ W, C. Roland 94-180, 7 July 1994. NUTZOTIN MOUNTAINS: rare on N-facing limestone talus slope in moist red silt clay, Cooper Pass, 1942 m, 62°17.16' N 142°31.44’ W, M. Cook 94353, 24 July 1994. WRANGELL MOUNTAINS: few in poplar wood- land, Nabesna River, 884 m, 62°12.63’ N 142°52.28’ W, M. Cook 96173B, 6 June 1996. This species is endemic to Alaska with a Pacific coastal distribution. There were two stations adjacent to the Park mapped in Hultén 1968, one at Chitina and one at Yakutat Bay (59°47.35'N 139°33.69'W, E.P. Walker 2416, 1914 (US)). The specimens cited above extend its range 111 km north into the Wrangell Mountains, 130 km north into the Nutzotin Mountains and 91 km south into the Chugach Mountains and document the northern extent of its range. Map 153. THE CANADIAN FIELD-NATURALIST Vol. 116 Astragalus nutzotinensis Rousseau, Nutzotin’s Milkvetch — CHUGACH MOUNTAINS: scattered in dry, loose marble gravel on SW-facing slope, ridge between Granite and Lime Creeks, 1829 m, 61°0.22’ N 141°51.1’ W, C. Roland 96-742, 28 July 1996; few on limestone outcrops and in patches of Dryas, ridge between Canyon, Pass and Tenas Creeks, 1682 m, 61°24.28' N 144°21.61' W, Cook & Losso 96327, 8 July 1996; locally common in sandy soil at contact between marble unit and darker crystalline rock on moderate W-facing slope, ridge between Canyon and Pass Creeks, 1463 m, 61°24.35' N 144°20.8' W, Roland & D'Auria 96-441, 9 July 1996. MENTASTA MOUNTAINS: scattered in limestone gravel on barren slope, ridge between Lost and Platinum Creeks at peak 5240, 1646 m, 62°34.58’ N 143°5.58' W, Cook & Roland 94334, 23 July 1994; rare in fine, needle-like argillite scree on steep, E- facing barren slope, ridge between Lost and Platinum Creeks at peak 5240, 1646 m, 62°34.58’ N 143°5.58' W, C. Roland 94-270, 23 July 1994; in runnels of soil in fine limestone scree, ridge between Little Jack and Trail Creeks, 1615 m, 62°36.05’ N 143°17.63' W, C. Roland 95-008, 6 June 1995; upper Trail Creek, 1341 m, 62°37.09’ N 143°16.06’ W, Potkin & Leggett 95-060A, 28 July 1995; scat- tered on moderate slope of calcareous gravel, Nabesna River at Totschunda Creek, 1280 m, 62°27.63' N 142°12.44' W, C. Roland 96-286, 24 June 1996. St. ELIAS MOUNTAINS: occasional in limestone scree below SE-facing cliffs, ridge between Dan & Copper Creeks, 1554 m, 61°21.56’ N 142°26.43' W, C. Roland 94-237, 14 July 1994; common in dry screes and ridges, Mt. Chitina, 2073 m, 60°57.74' N 141°17.33' W, C. Roland 95- 222, 16 July 1995. WRANGELL MOUNTAINS: moist sand and cobbles, confluence of Copper Creek and Kotsina River, 610 m, 61°42.34’ N 144°3.6’ W, C.R. Meyers 84-125, 27 July 1984; adjacent to limestone outcrop on gentle E-facing slope of saddle, Alice Peak, 1600 m, 61°39.8’ N 144°6.63' W, C.R. Meyers 84-144, 1 August 1984; rare in barren cobbles of river bar on W bank of Nabesna River at Monte Cristo Creek, 853 m, 62°13.71' N 142°54.14’ W, C. Roland 96-150, 18 June 1996; E-facing shaley scree, Nikolai Mine, 1695 m, 61°27’ N 142°39’ W, Batten & Barker 96-087, 24 July 1996; rare in loose gravel on barren SW-facing slope, volcanic ridge 6.4 km E of Snyder Peak, 1524 m, 62°4.47’ N 144°30.51' W, C. Roland 96-381, 5 July 1996; scattered in sandy exposed area on ridge above cliffs, plateau between Long Glacier and the Cheshnina River, 1399 m, 61°48.04’ N 144°6.27' W, M. Cook 96708, 8 August » 1996; occasional in moist sandy gravel on an unsta- ble slope, plateau between Long Glacier and the Cheshnina River, 1399 m, 61°48.04’ N 144°6.27' W, M. Cook 96710, 8 August 1996. This Alaska-Yukon endemic with an arctic-alpine distri- bution was known from seven localities in the northern 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 156. Lupinus kuschei 23 276 St. Elias and Wrangell Mountains: Guerin Glacier (61°37.42'N 141°4.38'W, D. F. Murray 2086, 4 August 1968 (ALA)); Sheep Glacier (61°42.48'N 141°38.59’'W, D. F. Murray 2241, 15 August 1968 (ALA)); Skolai Pass (61°37.25'N 141°58.77'W, D. F. Murray 711 (CAN) (Murray 1968); R. Scott 1631, 1746, 23 June 1967 (MICH) (Scott 1968)); Chitistone Pass (61°36.91'N 141°54.78'W, R. Scott 1631, 23 June 1967 (ALA) (Scott 1968); D. F. Murray 711, | July 1967 (ALA) (Murray 1968)); upper Chitina River (61°1.57'N 141°37.63’W (Hultén 1968)); Nugget Creek (61°37.51'N 143°43.63'W, E. Muphy s.n., 1978 (ALA)), and Nabesna (62°25.17'N 143°3.11'W (Hultén 1968)). Our collections extend its range into the western Wrangell, Mentasta and Chugach Mountains. It is considered rare in the Yukon Territory by Douglas et al. (1981). Map 154. Astragalus williamsii Rydb., Williams’ Milkvetch — MENTASTA MOUNTAINS: rare in open white spruce/poplar forest at toe of slope, Totschunda Creek, 1280 m, 62°27.63' N 142°12.44’ W, C. Roland 96-263, 24 June 1996; abundant at margin of floodplain white spruce forest, Nabesna River, 488 m, 62°47.9' N 142°9.97' W, C. Roland 96-301, 25 June 1996. This Alaska- Yukon endemic has a limited distribution in the state. The collections cited above extend its range 70 km south into the Mentasta Mountains from a station along the Slana-Tok Highway (Clearwater Creek, 62°59.38'N 143°20.35'W, Anderson & Brown 9224 (S) (Hultén 1941- 1950)). Map 155. Lupinus kuschei Eastw., Yukon Lupine — MENTASTA MOUNTAINS: occasional in sandy areas of gravel bar, Totschunda Creek floodplain, 725 m, 62°26.94' N 42 °40.8' W, C. Roland 96-255, 23 June 1996. WRANGELL MOUNTAINS: gravel roadside, km 2.4 Nabesna Road, 655 m, 62°41.84’ N 143°55.1’ W, M. Duffy 91014, 22 June 1991; sparsely vegetated dry forb-graminoid herbaceous meadow, Sanford Dunes, 686 m, 62°1.59’ N 145°0.12’ W, M. Duffy 92200, 10 July 1992. This Alaska- Yukon endemic with a boreal montane dis- tribution is rare in Alaska (G3 S2) and of limited distribu- tion in the Yukon Territory (Cody 1996). The specimens cited above extend its range 195 km into the Wrangell Mountains and 65 km into the Mentasta Mountains from a station along the Alaska Highway at mile 1257 (Lakeview Campground, 62°50.0’ N 142°00.0' W, H. Schmuck s.n., 16 August 1962 (ALA)). Map 156. Oxytropis campestris (L.) DC. ssp. jordalii (Pors.) Hult., Late Yellow Locoweed — MENTASTA MOUNTAINS: common in exposed, rocky site on SE- facing slope, Soda Lake, 1173 m, 62°32.36’ N 142°53.98' W, C. Roland 94-246B, 21 July 1994, M. Cook 94319, 22 July 1994 (ALA); common in exposed, rocky site on SE-facing slope, Totschunda Creek, 1280 m, 62°27.63’ N 142°12.44' W, C. Roland 96-272, 24 June 1996. WRANGELL MOUNTAINS: dry rocky soil on ridge crest, Nikolai Mine, 1695 m, 61°27’ N 142°39' W, Batten & Barker 96-061, 24 July 1996. THE CANADIAN FIELD-NATURALIST Vol. 116 This species is amphiberingean with an arctic-alpine dis- tribution. The collections cited above extend its range 314 km south into the Mentasta Mountains from a station in the Charley River Quad (vic. VABM Nation, 65°16.0' N 141°08.0' W, Lipkin & Cook s.n., 14 June 1993 (ALA)) and 277 km northwest of a station near Skagway (Hultén 1968). Map 157. Oxytropis huddelsonii Pors., Huddelson’s Locoweed — CHUGACH MOUNTAINS: solifluction soil, mesic tundra, Juniper Island, Granite Range, 1346 m, 60°36.17’ N 142°14.96' W, C. Roland 94-216, 12 July 1994; scattered in gravel, SW-facing slope, Granite Creek, 1829 m, 61°0.22’ N 141°51.1’ W, C. Roland 96-744, 28 July 1996; boulder field, in mat of Rhacomitrium, E. Fork Kiagna River, Granite Range, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-321, 29 July 1996 (ALA). MENTASTA MOUNTAINS: Soda Lake, 1173 m, 62°32.36’ N 142°53.98' W, C. Roland 94-259C, 22 July 1994; scattered in calcareous gravel, Totschunda Creek, 1280 m, 62°27.63’ N 142°12.44' W, C. Roland 96- 287, 24 June 1996; rare in dry, gravelly tundra, Devil’s Mountain, 1530 m, 62°25.62'’ N 142°53.93' W, C. Roland 96-360D, 3 July 1996. NUTZOTIN MOUNTAINS: moist tundra, NW-facing slope, 4 km E of Wiki Peak, 1585 m, 61°55.98' N 141°8.02’ W, C. Roland 94-119B, 23 June 1994; occasional on steep S-facing scree slope, Horsfeld Creek, 1768 m, 62°2.88' N 141°13.18' W, M. Cook 94146, 24 June 1994; scattered in gravel areas, S-facing alpine meadow, Carden Hills, 1311 m, 62°18.44’ N 141°11.55' W, C. Roland 94-140B, 26 June 1994; common in dry graminoid tundra, W-facing slope, Rock Lake, 1119 m, 61°48.7’ N 141°16.57’ W, C. Roland 96-032, 6 June 1996; occasional in tundra and dry gravels, Wiki Creek, 1411 m, 61°54.46’ N 141°10.68’ W, C. Roland 96-061, 6 June 1996. WRANGELL MOunrTAINS: N-facing scree slope, Lime Creek, 1707 m, 61°53.27’ N 141°37.05’ W, M. Cook 92112, 7 July 1992; basalt rubble, Cone Ridge, 2073 m, 62°8.21' N 143°18.47' W, M. Cook 94401, 25 July 1994; common in mesic tundra, Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85’ W, C. Roland 95-233, 18 July 1995; Dryas dwarf scrub tundra, Lakes Plateau, 1890 m, 62°4.4' N 143°23.5' W, Potkin & Leggett 95-113, 30 July 1995; occasional in morainal deposits, Ruddy Mountain, 1615 m, 62°4.61' N 144°46.39' W, C. Roland 96-329, 2 July 1996; dry tundra, Nikolai Pass, 1280 m, 61°26’ N 142°40' W, Batten & Barker 96-013, 23 July 1996 (ALA); dry tundra, Nikolai Pass, 1280 m, 61°26’ N 142°40’ W, Batten & Barker 96-049, 23 July 1996; common in tundra, Grotto Creek, 1847 m, 61°30.56’ ~N 142°24.79' W, C. Roland 96-647, 23 July 1996; moist scree, VABM Sentinel, W of Nizina Glacier, 1829 m, 61°39’ N 142°32’ W, Batten & Barker 96- 117, 24 July 1996 (ALA); scattered in volcanic sand, Cheshnina Plateau, 1399 m, 61°48.04’ N 144°6.27’ W, M. Cook 96712, 5 August 1996. 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 160. Callitriche anceps yd Se 278 This Alaska- Yukon endemic with a cordilleran distribu- tion had been collected at four localities in the Wrangell-St. Elias Mountains: Russell Glacier (61°41.93'N 141°45.36'W, D. F. Murray 2168, 11 August 1968 (ALA)); Guerin Glacier (61°37.42'N 141°4.38’W, D. F. Murray 2037, 3 August 1968 (ALA)); Chitistone Pass (61°37' N 141°58’ W, D. F. Murray 712 (CAN) (Murray 1968)); and Chetaslina River headwaters (61°59.36’N 144°20.24'W, R. Saltmarch PA21, 11 July 1978 (ALA)). The specimens cited above extend its range 114 km south into the Bagley Icefield, 71 km south into the Chugach Mountains, 96 km north into the Mentasta Mountains and 89 km north into the Nutzotin Mountains. This species is considered rare in Alaska (G3 S2S3). Map 158. Oxytropis scammaniana Hultén, Scamman’s Oxy- trope — NUTZOTIN MOUNTAINS: few on S-facing rub- ble slope, Wiki Creek, 1411 m, 61°54.46’ N 141°10.68’ W, M. Cook 96095, 7 June 1996; Dryas dwarf scrub tundra, Gold Hill, 1615 m, 62°6.88’ N 141°54.12' W, M. Cook 89930, C. Roland 96-071, 6 July 1989; scattered in gravel on S-facing subalpine meadow, Carden Hills, 1311 m, 62°18.44’ N 141°11.55’ W, C. Roland 94-139, 26 June 1994. WRANGELL MOUNTAINS: Dryas tundra, Lakes Plateau, 1890 m, 62°4.4' N 143°23.5' W, Potkin & Leggett 95-138, 31 July 1995; occasional in bare mineral soil, Cooper Pass, 1942 m, 62°17.16’ N 142°31.44’ W, M. Cook 94356, 24 July 1994; alpine tundra, Nabesna Glacier, 1768 m, 61°56.6’ N 143° W, M. Duffy 92119, 26 June 1992; dry rocky soil on ridge, Nikolai Mine, 1695 m, 61°27’ N 142°39’ W, Batten & Barker 96-062, 24 July 1996; occasional in morainal deposits, Ruddy Mountains, 1615 m, 62°4.61' N 144°46.39’ W, C. Roland 96-330, 2 July 1996; scattered in mesic tundra, Grotto Creek, 1661 m, 61°30.56’ N 142°24.79' W, C. Roland 96-656, 23 July 1996; scattered in crevices of basalt on ledge, Black Mountain, 1481 m, 62°20.85’ N 143°44.9’ W, M. Cook 95154, 2 July 1995; Dryas tundra, Upper Sanford River, 62°2.76' N 144°47.91' W, M. Duffy 91022, 23 June 1991. St. ELIAS MOUNTAINS: com- mon in N-facing tundra, Lime Butte, 1554 m, 61°21.56' N 142°26.43’ W, C. Roland 94-232A, 13 July 1994. This species is an Alaska-Yukon endemic with an arc- tic-alpine distribution. It had been collected at four locali- ties in the Wrangell-St. Elias Mountains: Chitistone Pass (Scott 1968); Skolai Pass (Murray 1968); Sheep Glacier (D.F. Murray 2260, 15 August 1968 (ALA)); and Russell Glacier (D.F. Murray 2187, 11 August 1968 (ALA)). The collections cited above extend its range 154 km west into the western Wrangell Mountains, 89 km north into the Nutzotin Mountains and connect the distribution 139 km to the northwest in the Alaska Range (Tangle Lakes, 63°03.0’ N 146°01.0’ W, G. Smith 2063, 21 August 1953 (ALA)) with the range 33 km to the east in the Yukon Territory (Cody 1996). Map 159. CALLITRICHACEAE Callitriche anceps Fern. (Callitriche heterophylla ssp. heterophylla Pursh.) — CHUGACH MOUNTAINS: THE CANADIAN FIELD-NATURALIST Vol. 116 beaver pond, Three Mile Canyon, 610 m, 60°58.37' N 144°8.44’ W, M. Duffy 91148, 19 August 1991. NUTZOTIN MOUNTAINS: submerged in shallow water of small remnant pond in kettle depression, Braye Lakes, 1097 m, 62°2’ N 141°9’ W, Parker & Gracz 6887, 11 August 1996. WRANGELL MOUNTAINS: sub- merged in slow moving creek, vic. Grizzly Lake, 62°13.92'N 143°22.17’'W, Cook & Allen 3559, 26 July 2000. This North American boreal montane aquatic plant is considered rare in the Yukon Territory (Douglas et al. 1981). The stations cited above extend its range 347 km east into the Chugach Mountains from a station on the Kenai Peninsula (Hultén 1968), 252 km south into the Wrangell Mountains and 338 km southeast into the Nutzotin Mountains from a collection in the Big Delta Quad (Birch Lake, 64°20.0' N 146°40.0’ W, C. Parker 2200, 29 Aug 1989 (ALA)). Map 160. Callitriche hermaphroditica L., Northern Water- starwort — NUTZOTIN MOUNTAINS: lake margin, 0.5 m deep, mixed freshwater herbaceous, Ptarmigan Lake, 1079 m, 61°51.51' N 141°10.78' W, J. Barnes 96-252, 8 August 1996. WRANGELL MOUNTAINS: scattered in 50 cm of water, rooted in rocky floor of small, shallow pond, Fish Creek, 1067 m, 62°16.92’ N 142°58.99' W, C. Roland 95-284, 29 July 1995. This circumpolar boreal montane species has a spotty distribution throughout its range. The specimens cited above extend its range 99 km south into the Nutzotin Mountains from a station along the Alaska Highway near the Yukon border (Nabesna Quad: Scottie Creek, 62°41.0’ N 141°05.0’ W, L. Vining 61, 27 August 1979 (ALA)) and 179 km southeast into the Wrangell Mountains from a collection at Tangle Lakes (Mt. Hayes Quad: 63°03.0' N 146°01.0' W, G. Smith 2090, 22 August 1953 (ALA)). Map 161. BALSAMINACEAE Impatiens noli-tangere L., Western Touch-Me-Not — CHUGACH MOUNTAINS: Rare in the shade of tall alder scrub, confluence of Bremner and Copper Rivers, 152 m, 61°0.94' N 144°40.76’ W, C. Roland 96-597, 17 July 1996. The locality cited above of this incompletely circumpo- lar widespread species, extends its range 87 km east into the Chugach Mountains from a station near Cordova (Hultén 1968). The closest station to the east is 550 km near Skagway (Hultén 1968). Map 162. VIOLACEAE Viola adunca J.E. Smith, Hook-spur Violet — ST. ELIAS MOUNTAINS: moist herbaceous patch among birch shrubs, Short River, 503 m, 61°5.35’ N 141°56.3' W, Parker & Duffy 6702, 6 August 1996. CHUGACH MOUNTAINS: occasional in moss of mead- ow, Nerelna Creek Plateau, 1372 m, 61°26.66’ N 144°17.69' W, M. Cook 96302, 7 July 1996; herba- ceous meadow, Upper Golconda, 1311 m, 61°2.8’ N 143°25.64’ W, L. A. & E. G. Viereck 11072, 7 July 1996; W-facing heath, Juniper Island, 1291 m, 60°36.24' N 142°21.69' W, C. Roland 94-218A, 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 279 161. Callitriche hermaphroditica 164. Viola biflora 280 12 July 1994; mesic tundra slope, Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96-205, 27 July 1996. This violet is North American with a boreal-montane distribution. The collections cited above extend its range 103 km south into the Chugach Mountains from a collec- tion on Bonanza Ridge in the Wrangell Mountains (Nordell & Schmitt 1977) and connect the range 356 km to the west in the Anchorage Quad (61°01.0' N 149°44.0' W, L.L. Hawkins s.n., 12 June 1976 (ALA)) with the range in the Yukon Territory 207 km to the east (Cody 1996). Map 163. Viola biflora L., Twoflower Violet — WRANGELL MOuNnNTAINS: E-facing rubble slope, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24’ W, Moran & Roland 95-34, 1 July 1995; few, in Rhacomitrium between boulders along ridge, Black Mountain, 1481 m, 62°20.85'’ N 143°44.9' W, Cook & Beck 95142, 1 July 1995; few under boulders on rubble slope, Fish Creek , 1067 m, 62°16.92’ N 142°58.99’ W, M. Cook 95320, 27 July 1995; abundant on east and W-facing morainal slopes, Grizzly Lake, 1108 m, 62°19.51'N 143°9.24'W, Allen & Cook 00-067, 25 July 2000. This violet has an incompletely circumpolar distribution throughout arctic-alpine regions and is considered rare in the Yukon Territory (Douglas et al. 1981). The four locali- ties cited above extend its range 151 km south into the Wrangell Mountains from the Alaska Range (Mt. Hayes Quad: 63°38.0' N 144°44.0' W, L.A. Spetzman 183, 25 June 1957 (ALA)) and 261 km east from the Talkeetna Mountains (62°31.0’ N 149°11.0’ W, E. Helmstetter 213- 79, 29 July 1979 (ALA)). Map 164. Viola selkirkii Pursh, Great-spurred Violet — WRANGELL MOUNTAINS: growing in alder-spruce stand, near Angle Station, Bonanza Ridge, 975 m, 61° 30.70’ 142° 49.85’, G. Frost s.n., 1 June 1998 (ALA). This boreal-montane violet is circumpolar with large gaps. The locality cited above from the Wrangell Mountains connects its range 180 km to the southwest in the Valdez Quad (Mile 14 Richardson Highway, I. W. Ailes 17 (ALA)) with collections 473 km to the southeast near Skagway (Hultén 1968) and 615 km to the northeast in the Yukon Territory (Cody 1996). It is considered rare in Alaska (G5? $3) and in the Yukon Territory (Douglas et al. 1981). Map 165. ONAGRACEAE Epilobium lactiflorum Haussk, White-Flower Willowherb — CHUGACH MOUNTAINS: forb herba- ceous meadow, Juniper Island, 1291 m, 60°36.24' N 142°21.69' W, M. Cook 94283, 13 July 1994. St. ELIAS MOUNTAINS: alpine mesic forb herbaceous slope, Blossom Island, 732 m, 59°59.97' N 140°5.1’ W, M. Cook s.n., 21 August 1987. WRANGELL MOUNTAINS: clumped in mesic dry creek bed, Nadina Glacier, 1768 m, 62°2.85' N 144°41' W, M. Cook 94420, 30 August 1994; in Aulocomnium moss in stream, Chetaslina Ridge, 1615 m, 61°56.51’ N 144°25.93' W, M. Cook 94494, 17 August 1994. THE CANADIAN FIELD-NATURALIST Vol. 116 This species is amphiatlantic with a boreal-montane dis- tribution. It was collected by O. Nordell and A. Schmitt on Bonanza Ridge in the southern Wrangell Mountains (Nordell and Schmitt 1977). The Wrangell Mountain col- lections cited above extend the range of this species 116 km into the Western Wrangells and the St. Elias Mountain col- lections connect the range 315 km to the west near Cordova (Schwan Glacier, 60°58.0’ N 145°00.0’ W, C. Parker 1766, 12 August 1986 (ALA)) with a station near Haines (Cody 1996). Map 166. Epilobium luteum Pursh, Yellow Willowherb — St. ELIAS MOUNTAINS: mesic mixed herbaceous alpine slope, Samovar Hills, 549 m, 60°8.11’ N 140°39.55’ W, M. Cook 87-143, 23 August 1987. This species is North American with a Pacific coastal distribution. The collection cited above connects the range 330 km to the west near Cordova (Hultén 1968) with a station at Yakutat 110 km to the east (Hultén 1968). Map 167. HIPPURIDACEAE Hippuris montana Ledeb., Mountain Marestail — GULF OF ALASKA: dwarf scrub/mixed forb alpine tundra, Icy Bay, 579 m, 60°5.83’ N 141°27.3’ W, K.A. Beck s.n., 10 August 1987; tall forb meadow, Chaix Hills, 1550 m, 60°3.18’ N 141°5.1’ W, M. Duffy 92213, 17 August 1992.) WRANGEER MOUNTAINS: mud along shore of pond, 0.8 km east of Lake 2910’, Copper River, 823 m, 62°26.35’ N 143°40.77' W, M. Cook 96744, 8 August 1996. This North American Pacific coastal species is known from one locality in the Yukon Territory and should be added to the list of rare plants for the Territory (Cody 1994, 1996). The specimens cited above from Icy Bay and Chaix Hills connect its coastal range 394 km to the west at Thum Bay (W.J. Eyerdam 3301, 3537, 31 July 1931 (UC, S) (Hultén 1946)) with a station 357 km to the east near Haines (Hultén 1968). The station cited above from the Wrangell Mountains extends the range of this species 297 km to the southeast from a station near Denali National Park (Hultén 1968) and is 298 km north of the coastal localities in the Park. Map 168. HALORAGACEAE Myriophyllum verticillatum L., Whorlleaf Watermil- foil — WRANGELL MOUNTAINS: | m deep water, bog 2.5 km NW of Billy Lake, 614 m, 61°27.99' N 143°56.38' W, C. Roland 94-323, 10 August 1994. This species is incompletely circumpolar and disjunct from temperate zones. It is rare in Alaska (GS S3) and the Yukon Territory (Douglas et al. 1981). The specimen cited above extends its range 238 km east from the Anchorage Quad (Otter, 61°17.53’ N 149°44.17’ W, Duffy & Tande 1019, 3 August 1994 (ALA)) and connects the range 363 km to the east near Haines Junction in the Yukon Territory (Cody 1996). Map 169. APIACEAE Cicuta maculata L. var. angustifolia Hook. (C. dou- glasii (DC.) Coult. & Rose), Spotted Water Hemlock — GULF OF ALASKA: wet graminoid meadow, Yahtse River mouth, Malaspina Forelands, 30 m, 59°50.22’ N 141°13.67' W, M. Duffy 92237, 19 August 1992. 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 168. Hippuris montana 281 —— —_— ss 282 This specimen connects the Pacific Coastal distribution of this North American species 186 km to the west at Cape St. Elias (59°56.0’ N 144°23.0' W, Cunningham & Stanford 228-78, 21 June 1978 (ALA)) with stations in the vicinity of Yakutat and the Ankow River 89 km to the east (F. Funston 71, 1892 (CAN, UC) (Coville and Funston 1896; Hultén 1946); C.V. Piper 42, 82, 1904 (US) (Hultén 1946)). This species is considered rare in the Yukon Territory (Douglas et al. 1981). Map 170. Ligusticum scoticum L. ssp. hultenii (Fern.) Calder & Taylor, Hultén’s Sea-lovage — GULF OF ALASKA: closed tall willow-alder scrub, 0.2 km N of Schooner Beach, Malaspina Forelands, 30 m, 59°46.52’ N 140°1.45’ W, K.A. Beck s.n., 9 August 1987. This species is amphiberingian with a North Pacific Coastal distribution. The specimen cited above connects its range 253 km to the west at Cape St. Elias (59°56.0' N 144°23.0' W, Cunningham & Stanford 198-78, 16 June 1978 (ALA)) with a station in the vicinity of Yakutat 30 km to the east (F. Funston 70, 1892 (US,UC); Trelease & Saunders 4530, 4531, 1899 (US) (Hultén 1946)). Map 171. Osmorhiza depauperata Phil., Bluntseed Sweetroot — CHUGACH MOUNTAINS: open cottonwood forest, Jackson Creek, 61°7.38’ N 144°53.17' W, M. Duffy 91133, 15 August 1991. This North American boreal-montane umbel is rare in the Yukon Territory (Douglas et al. 1981). The locality cited above represents the northern extent of its range and connects the range 128 km to the west near Valdez (Hultén 1968) with the range 582 km to the southeast near Gustavus (Hultén 1968). Map 172. ERICACEAE Cassiope mertensiana (Bong.) D. Don, White Mountain Heather — CHUGACH MOUNTAINS: moun- tain heath tundra on scoured bedrock, ridge east of Falls Creek falls, 1097 m, 61°9.2’ N 144°22.31' W, M. Duffy 91116, 12 August 1991. This species is North American with a Pacific coastal distribution. The locality cited above connects a station 939 km to the southwest in the Chignick Quad (56°18.0' N 158°24.0’ W, J. Dodge s.n., 24 July 1985 (ALA)) with sta- tions 489 km to the southeast near Haines Junction and Skagway (Hultén 1968). Cody et al. (1998) reported this species new to the Yukon in the southeast corner and sug- gested that it be added to the list of rare species in the Yukon Territory. Map 173. Cladothamnus pyrolaeflorus Bong. (Elliottia pyro- liflorus (Bong.) S.W. Brim & P.F. Stevens), Copper Flower — St. ELIAS MOUNTAINS: heather/mesic forb alpine tundra, Floral Pass, 549 m, 59°57.91’ N 139°57.76' W, M. Cook 87-71, 21 August 1987. This specimen connects the range of this North American Pacific Coastal species 322 km to the west near Cordova (B. J. Neiland 1343, 2 July 1965 (ALA)) with sta- tions 249 km to the east near Haines in the Mt. Fairweather Quad (Lituya Bay, 58°40.0’ N 137°36.0' W, S. Doman 85-46, 16 June 1985 (ALA)). Map 174. Phyllodoce glanduliflora (Hook.) Cov. (P. aleutica (Spreng.) Heller ssp. glanduliflora (Hook.) Hult.), THE CANADIAN FIELD-NATURALIST. Vol. 116 Aleutian Mountain Heather — CHUGACH MOUNTAINS: mountain heath tundra, Falls Creek, 1097 m, 61°9.2' N 144°22.31’ W, M. Duffy 91115, 12 August 1991; sandy dry soil in depression sur- rounded by Empetrum nigrum-Salix arctica heath, Juniper Island, 1346 m, 60°36.17’ N 142°14.96’ W, M. Cook 94232, 10 July 1994; co-dominant in mesic heath, 1291 m, 60°36.24' N 142°21.69’ W, C. Roland 94-217, 12 July 1994. St. ELIAS MOUNTAINS: dwarf scrub/mixed forb herbaceous alpine tundra, Guyot Glacier, 579 m, 60°5.83’ N 141°27.3’ W, K. Beck s.n., 10 August 1987; alpine tundra heath, Samovar Hills, 549 m, 60°8.11' N 140°39.55’ W, M. Cook 87-154, 23 August 1987. WRANGELL MOUNTAINS: dominant on NE-facing slope, Mt. Drum, 1433 m, 62°8.83’ N 144°30.18' W, M. Cook 94056, 11 June 1994. This species is North American with a cordilleran distri- bution. The localities cited above extend its range 161 km northeast into the Wrangell Mountains and 232 km east into the Chugach Mountains from a station near Valdez (Hultén 1968). The specimens from the southern St. Elias Mountains connect the range 113 km to the east at Yakutat (Hultén 1968). Map 175. PRIMULACEAE Douglasia alaskana (Coville & Standley ex Hultén) S. Kelso (Androsace alaskana Cov. & Standl.), Alaskan Douglasia — CHUGACH MOUNTAINS: few on old mining road, Five Mile Creek, 1463 m, 61°32.58' N 143°58' W, M. Cook 93463, 27 August 1993; few on talus slope and Dryas tundra, Hundell Creek, 1585 m, 61°36.75’ N 144°42.2' W, M. Cook 93467, 28 August 1993. WRANGELL MOUNTAINS: few along ridge at head of Elliot Creek, 1926 m, 61°38.99'’ N 144°3.51’ W, M. Cook 96468, 16 July 1996. This primrose is an Alaska-Yukon endemic with a Pacific Coastal distribution that is considered rare in Alaska (G2G3 S2S) and in the Yukon (Douglas et al. 1981). The collections cited above extend its range 203 km east into the Chugach and Wrangell Mountains from collections in the Anchorage Quad (Lower Snowhawk, 61°11.75’ N 149°33.25' W, M. Duffy 795, 19 July 1994 (ALA)) and connect the range 275 km to the east in the Yukon Territory (Cody 1996). Map 176. Douglasia arctica Hook., Mackenzie River Dwarf Primrose — ST. ELIAS MOUNTAINS: rare in rocky sites on steep, N-facing slopes, Mt. Natazhat, 1716 m, 61°35.38’ N 141°1.83’ W, C. Roland 95-066, 19 June 1995. This primrose is an Alaska- Yukon endemic with an arc- tic-alpine distribution and is rare in Alaska (G3 S283). The collection cited above extends its range 297 km south into the St. Elias Mountains from a station near Chicken (Hultén 1968) and 296 km east from a station near Valdez (Hultén 1968). Map 177. Douglasia gormanii Const., Gorman’s Dwarf Primrose — MENTASTA MOUNTAINS: occasional in moist soil and rubble on NE-facing debris flow, Trail 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 171. Ligusticum scoticum ssp. hultenii 172. Ozmorhiza depauperata 283 ———_—- SO -—- +--+ ~~ 284 THE CANADIAN FIELD-NATURALIST Vol. 116 176. Douglasia alaskana 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 180. Primula egaliksensis 285 286 Creekf 1615 m, 62°36:05"" N 143717363" WTC: Roland 95-014B, 6 June 1995; Upper Trail Creek, 1341 m, 62°37.09’ N 143°16.06’ W, Potkin & Leggett 95-054, 28 July 1995; rare near top of steep talus slope, occasional on N-facing limestone rubble slope, headwaters of Lost Creek, 1722 m, 62°36.45' N 143°12.03’ W, K.A. Beck 95-222, 4 July 1995; M. Cook 95171, 7 July 1995; occasional in fine gravel, ridge above Totschunda Creek, 1280 m, 62°27.63' N 142°12.44’ W, C. Roland 96-293, 24 June 1996; scattered in rubble chute on NE-facing slope, Devil's Mountain, 1530 m, 62°25.62' N 142°53.93’ W, C. Roland 96-350, 3 July 1996. NUTZOTIN MOUNTAINS: rare on upper slope, Carl Creek, 1920 m, 62°3.52’ N 141°36.27' W, M. Cook 94095, 20 June 1994; patchy on rock outcrops on northwest side of ridge, Wiki Peak, 1433 m, 61°53.9' N 141°9.51' W, M. Cook 94123, 22 June 1994; occasional in moist tundra vegetation among cobble-sized rocks, slope 4 km east of Wiki Peak, 1585 m, 61°55.98’ N 141°8.02’ W, C. Roland 94-118B, 23 June 1994; occasional on SW-facing scree slope, Horsfeld Creek, 1768 m, 62°2.88' N 141°13.18' W, C. Roland 94-129B, 24 June 1994; rare on southeast trending limestone ridces Bawttolf Creek, 1707 m, 62°92,13"-> N 141°14.51’ W, M. Cook 94190, 28 June 1994; occa- sional on ridge growing in Rhacomitrium, Klein Creek, 1747 m, 62°2.29’ N 141°19.88’ W, M. Cook 95017, 15 June 1995; few on ridge in rocky dwarf scrub tundra, Rock Lake, 1119 m, 61°48.7' N 141°16.57' W, M. Cook 96028, 6 June 1996; scat- tered mid-slope in dry sandy gravel, 1411 m, 61°54.46’ N 141°10.68' W, M. Cook 96060, 6 June 1996; common but scattered on gravel slope, Wiki Basin, 1411 m, 61°54.46’ N 141°10.68' W, C. Roland 96-067, 6 June 1996; rare on well vegetated SE-facing alpine colluvium, Horsfeld Creek valley, 1128 m, 62°2’ N 141°11' W, Parker & Gracz 6915, 13 August 1996; bluff east of lodge, Ptarmigan Lake, 1079 m, 61°53’ N 141°8.8' W, Duffy & Barnes 96- 259, 8 August 1996; scattered on NW-SE trending ridge amongst Dryas stripes, pass between Cabin and Wiki Creeks, 1585 m, 61°53.64’ N 141°11.4’ W, M. Cook 3172, 14 June 1998 (ALA); steep S-facing rhyolite scree knob, W side of Wiki Creek, 1524 m, 61°54.77' N 141°11.05' W, M. Cook 3181, 15 July 1998 (ALA); scattered in gravel patches amongst Dryas stringers, Ophir Creek, 1463 m, 61°55.7' N 141°31.07’ W, M. Cook 3191, 16 July 1998 (ALA); scattered in sparsely vegetated basalt scree, ridge S of Sonja Creek, 1494 m, 61°57.14’ N 141°21.43’ W, M. Cook 3194, 16 July 1998 (ALA). WRANGELL MOUNTAINS: alpine tundra, Nabesna Glacier, 1768 m, 61°56.6' N 143° W, M. Duffy 92115, 26 June 1992; N-facing scree slope, ridge between Lime Creek and Solo Flats, 1707 m, 61°50.85’ N 141°47.12' W, M. Cook 92110, 7 Suly 1992; occa- sional on steep gravel slope, northeast slope of Mt. THE CANADIAN FIELD-NATURALIST Vol. 116 Drum, 1433 m, 62°8.83’ N 144°30.18’ W, C. Roland 94-055, 11 June 1994; occasional on slope of exposed, stony alluvium, Mt. Sanford cinder cone, 1722 m, 62°23.4’ N 144°15.9’ W, C. Roland 94-002, 6 June 1994; rare on steep W-facing gravel slopes, Cone Ridge, 2073 m, 62°8.21’ N 143°18.47’ W, C. Roland 94-318B, 27 July 1994; sparsely vegetated disturbed S-facing rubble, Chetaslina Plateau, 1615 m, 61°56.51’ N 144°25.93’ W, C. Roland 94-351, 17 August 1994; occasional in stony, frost boiled poly- gon tundra, Jaegar Mesa, 1893 m, 62°15.9’' N 143°1.24’ W, C. Roland 95-115, 1 July 1995; occa- sional in crevices of basalt rock ledges on ridge, Black Mountain, 1481 m, 62°20.85’ N 143°44.9’ W, M. Cook 95153, 7 July 1995; southwest slope of Mt. Drum, between Nadina Glacier and Klawasi River, occasional in stony areas in sparse tundra, 1615 m, 62°4.61’ N 144°46.39’' W, C. Roland 96-328, 2 July 1996. This species, an Alaska-Yukon endemic with a cordilleran distribution, is rare in Alaska (G3 $3). It had been collected at four localities in the Wrangell-St. Elias Mountains: Guerin Glacier (61°37.42'N 141°4.38’W, D. F. Murray 2046, 3 August 1968 (ALA); Sheep Glacier (61°42.48'N 141°38.59'W, D. F. Murray 2278, 17 August 1968 (ALA)); Russell Glacier (61°41.93'N 141°45.36'W, D. F. Murray 2169, 11 August 1968 (ALA)) and Chitistone Pass (61°37' N 142°3' W, R. Scott 1974, 94G6 190s (MICH) (Scott 1968) and D. F. Murray 722 (CAN) (Murray 1968)). The new collections cited above extend its range 151 km into the northern and western Wrangells, 128 km northwest into the Mentasta Mountains and 90 km north into the Nutzotin Mountains. Map 178. Primula cuneifolia Ledeb. ssp. saxifragifolia (Lehm.) Sm. & Forrest, Pixie-Eyes — CHUGACH MOUNTAINS: lush meadow on the margin of a seep in E-facing tun- dra, Juniper Island, 1346 m, 60°36.17’ N 142°14.96’ W, C. Roland 94-194, 10 July 1994; scattered adja- cent to creek in dwarf willow-crowberry heath, plateau between Grant Creek, Tebay River and Hanagita River, 1250 m, 61°17.65' N 143°56.34’ W, M. Cook 96355, 7 July 1996; several plants along the edge of a small stream, Upper Golconda Creek, 1280 m, 61°3’ N 143°24.45’ W, L. A. & E.G. Viereck 11054, 7 July 1996. GULF OF ALASKA: ericaceous dwarf scrub alpine tundra, tall forb herbaceous, 3.5 km NW of Hanna Lake, Robinson Mts., 610 m, 60°9.95' N 143°6.8' W, M. Duffy 9256, 13 June 1992; ericaceous dwarf scrub alpine tundra & scree slope, nunatak between peaks 2710 and 2425, Robinson Mountains, 646 m, 60°9.9' N 143°10’ W, M. Cook 9210, 13 June 1992. WRANGELL MOUNTAINS: scat- tered in mesic Dryas tundra lining an intermittent _stream channel on gentle SE-facing slope, Upper Dadina River, 1344 m, 62°3.95’ N 144°25.18’ W, C. Roland 96-312, 1 July 1996; scattered throughout basin in moss and organic soil of ericaceous tundra, upper Dadina River, 1344 m, 62°3.95' N 144°25.18’ W, M. Cook 96268, 7 July 1996. 2002 CooK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 287 This amphiberingian species has a north Pacific coastal distribution. The specimens cited above extend the range 135 km south into the Wrangell Mountains from a collec- tion in the Alaska Range (Maclaren Summit, 63°05.0’ N 146°23.0’' W, D. Rhode 81/P6-85, 20 June 1981 (ALA)), 200 km east into the Granite Range from collections at Thompson Pass (61°7.81'N 145°42.11'W, J.P. Anderson 1910, July 1935 (Hultén 1947)) and connect the distribu- tion 368 km to the southeast near Haines (Hultén 1968). This species may have been collected by the Alaska Boundary Survey in 1912 at high elevations in the Wrangell Mountains. This collection was described as “Wrangell at high altitudes” by Hultén (1947) but mapped within the Eastern Pacific Coastal district. However, there are no records that the Alaska Boundary Survey visited Wrangell Island and they did travel through Skolai Pass in 1912 (Green 1982; personal communication with Geoff Bleakley, National Park Service historian, 10 October, 2000). Map 179. Primula egaliksensis Wormsk., Greenland Primrose — MENTASTA MOUNTAINS: along stream, Soda Lake, 1494 m, 62°31.2’ N 142°54.03’ W, Moran & Roland s.n., 5 July 1995; rare in Dryas tundra on limestone, Soda Lake, 1173 m, 62°32.36’ N 142°53.98’ W, C. Roland 94-255B, 22 July 1994. WRANGELL MOUNTAINS: lakeshore gravel, Jack Lake, 975 m, 62°31.2' N 143°18.54’ W, M. Cook 3021, 20 June 1997. This North America arctic-alpine primrose is rare in the southern part of the Yukon Territory (Cody 1996). The localities cited above extend its range 167 km to the south into the Mentasta Mountains from collections in the Alaska Range (Fielding Lake, 63°10.0’ N 145°41.0’ W, G. Smith 2614, 8 July 1955 (ALA)). Map 180. GENTIANACEAE Fauria cristi-galli (Menzies) Makino, Deer Cabbage — GULF OF ALASKA BASIN: mixed forb meadow, Independence Creek, Icy Bay, 427 m, 60°0.76’ N 141°27.1' W, K. Beck s.n., 12 August 1987; erica- ceous shrub/graminoid herbaceous bog, Cape Yakataga, 50 m, 60°2.7' N 142°8’ W, M. Cook 9201, 10 June 1992; tall forb herbaceous alpine tun- dra, Robinson Mountains, 671 m, 60°5.64’ N 142°17.17' W, M. Duffy 9234, 12 June 1992. This species is North American with a North Pacific coastal distribution. The collections cited above connect the range 262 km to the west near Valdez (Hultén 1968) with collection 167 km to the southeast near Pelican (Hultén 1968). Map 181. Gentiana douglasiana Bong., Swamp Gentian — CHUGACH MOUNTAINS: subarctic lowland sedge moss bog meadow, Martin River Glacier, 91 m, 60°14.65’ N 144°1.15' W, M. Duffy 92265, 3 September 1992. This species is North American with a Pacific coastal distribution. The specimen cited above connects the range 99 km to the west near Cordova (Hultén 1968) with a local- ity 160 km to the east at Icy Bay (Hultén 1968). Map 182. Gentiana platypetala Griseb., Broadpetal Gentian — ST. ELIAS MOUNTAINS: forb meadow, Moore Nunatak, 610 m, 60°4.67'’ N 140°54.93’ W, M. Duffy 92221, 18 August 1992. This gentian is an Alaskan endemic with a Pacific coastal distribution. The collection cited above connects the range 305 km to the west at Hawkins Island (Hultén 1968) with the range 103 km to the east at Yakutat (Hultén 1968). Map 183. Gentianella tenella (Rottb.) Borner, Dane’s Dwarf Gentian — WRANGELL MOUNTAINS: abundant in Sibbaldia procumbens dominated forb herbaceous vegetation on S/SW-facing snowbed gully, ridge between Chichokna and Chetaslina Rivers, 1615 m, 61°56.51' N 144°25.93' W, C. Roland 94-348, 94- 350, 17 August 1994. This circumpolar arctic-alpine species was known from the Wrangell Mountains (Chitistone Pass, 61°37’ N 142°3' W, R. Scott s.n., 1967 (MICH) (Scott 1968)). The new locality cited above extends the range 138 km to the west in the Wrangell Mountains. These populations connect the distribution 247 km to the west in the Anchorage Quad (Eklutna Valley, 61°24.0’ N 149°04.0’ W, L. Marvin 1800, 13 August 1984 (ALA)) with the distribution in the Yukon Territory 217 km to the east (Cody 1996). Map 184. Swertia perennis L., Star Gentian — CHUGACH MOUNTAINS: patchy in mesic Aulocomnium palustre, under birch scrub, Sangaine Creek, 732 m, 61°13.13’ N 143°56.26' W, M. Cook 96392, 7 July 1996. GULF OF ALASKA: Malaspina Forelands, 61 m, K.A. Beck s.n., August 1987. This species is incompletely circumpolar and has a bore- al-montane distribution. It was known previously in the vicinity of the Park at Yakutat (Stair & Pennell s.n., 13 July 1945 (PH) (Stair and Pennell 1946; Hultén 1941-1950)); the 141st meridian north of Mt. St. Elias (D.W. Eaton 1], (US) (Hultén 1941-1950)) and Indian Creek (W.L. Poto 151, 16 August 1902 (US) (Poto 1902*, Hultén 1941-1950)). The new collection cited above extends its range 111 km north- east into the Chugach Mountains from collections near Cordova (Hultén 1968) and connects the range approxi- mately 162 km to the southeast at Yakutat. Map 185. POLEMONIACEAE Collomia linearis Nutt., Narrowleaf Mountain- trumpet — WRANGELL MOUNTAINS: gravel bar, McCarthy Creek, 61°24.51' N 142°50.27' W, K.A. Beck s.n., 6 August 1988. This species is native to western North America but introduced north to the District of Mackenzie, Yukon Territory, Alaska and eastward (Cody 1996). It is known from a few widely disjunct localities in Alaska. The speci- men cited above is 394 km south of a collection at Eagle Bluff (64°47.0’ N 141°12.0' W, M. Moldenhauer Y234, 13 July 1982 (ALA)), 379 km east of a collection near Anchorage (Hultén 1968) and 953 km northwest of a col- lection in the Ketchikan Quad (Hyder, 55°55.0' N 130°00.0' W, J.P. Anderson 5430, 28 June 1939 (ALA)). Map 186. Phlox hoodii Richards., Spiny Phlox — MENTASTA MOUNTAINS: locally abundant in silty soil on steep, S-facing slopes, Devil's Mountain, 942 m, 62°24.95' N 142°54.86' W, C. Roland 96-235, M. Cook 96239, 22 June 1996. This North America prairie species is disjunct from the temperate zone in Alaska and southwestern and central 288 THE CANADIAN FIELD-NATURALIST Vol. 116 181. Fauria crista-galli 184. Gentianella tenella 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA Phlox richardsonii 289 290 Yukon Territory (Cody 1996). It is considered rare in Alaska (G5 S182). The collection cited above from the Mentasta Mountains extends its range 290 km south from collections at Eagle Bluff (64°48.0’ N 141°12.0’ W, Battten & Dawe 78-253, 14 July 1978 (ALA)) and connects the distribution 194 km to the east in the Yukon Territory (Cody 1996). Map 187. Phlox richardsonii Hook. (Phlox sibirica L. ssp. richardsonii (Hook.) Hult.), Siberian Phlox — MENTASTA MOUNTAINS: localized on S-facing green- stone outcrop, above Soda Creek, 1173 m, 62°32.36’ N 142°53.98’ W, C. Roland 94-260, 22 July 1994; occasional in rock crevices, SE-facing slope, Devil's Mountain, 1530 m, 62°25.62’ N 142°53.93’ W, C. Roland 96-355, 3 July 1996. NUTZOTIN MOUNTAINS: S-facing scree, 2 km W of Ptarmigan Creek, 1707 m, 61°55.65' N 141°6.5" WM. Duffy 92179.. 7 -Suly 1992; S-facing scree slope, 2.4 km W of Ptarmigan Creek, 1494 m, 61°55.65’ N 141°6.5’ W, M. Cook 92478, 9 July 1992; locally abundant in gravel on SW-facing slope, Upper Wiki Creek, 1433 m, 61°53.9’ N 141°9.51' W, C. Roland 94-116A, 22 June 1994; S-facing talus slope, ridge 1.6 km E of Rocker & Ptarmigan Creek confluence, 1433 m, 61°55.22’ N 141°2.75' W, Cook & Roland 94143, 23 June 1994; locally common in rock crevices, Klein Creek, 1747 m, 62°2.29’ N 141°19.88' W, C. Roland 95-031, 15 June 1995; scattered in orange rhyolite, volcanic plateau between Bryan and Willow Creeks, 1829 m, 61°58.97' N 141°50.03’ W, M. Cook 95051, 18 June 1995; scattered in gravel on S-facing slopes, Rock Lake, 1119 m, 61°48.7’ N 141°16.57’ W, ™. Cook 96040, C. Roland 96-052, 6 June 1996; com- mon in barren, sand and gravel, Wiki Creek, 1411 m, 61°54.46' N 141°10.68’ W, C. Roland 96-068, 6 June 1996; scattered in basalt scree, Sonja Creek, 1494 m, 61°57.29’ N 141°21.29' W, M. Cook 3192, 16 July 1998 (ALA). WRANGELL MOUNTAINS: N-fac- ing scree slope, Lime Creek, 1707 m, 61°47.2' N 141°49.06’ W, M. Cook 92108, 9 July 1992; Dryas sedge tundra, NW slope of Mt. Sanford, 1295 m, 62°21.17' N 144°24.52’ W, M. Cook 93119, 9 June 1993; S-facing volcanic slope, Capital Mountain, 1052 m, 62°31.61' N 144°12.38' W, C. Roland 94- 015, 7 June 1994; scattered on S-facing slope, Mt. Sanford, 1615 m, 62°13.59’ N 144°26.06’ W, C. Roland 94-041, 9 June 1994; scattered in rock out- crop crevices on bluff, Nabesna River, 1106 m, 62°15.05' N 142°54.68’ W, M. Cook 96195, 6 June 1996; occasional on S-facing scree slope in moist bare sandy soil, Carl Creek, 1920 m, 62°3.52' N 141°36.27' W, M. Cook 95004, 14 June 1994. This Alaska-Yukon endemic with an arctic-alpine distri- bution is rare in Alaska (G4T2T3Q S27). It was known from two localities in Alaska, one in the Yukon Territory, the Arctic Coast of the District of Mackenzie and Banks Island (Cody 1996; Porsild 1980). The stations cited above extend its range 345 km south into the Mentasta, Nutzotin and Wrangell Mountains and connects the range in the Yukon Territory, 102 km to the southeast (Cody 1996). Map 188. THE CANADIAN FIELD-NATURALIST Vol. 116 HYDROPHYLLACEAE Phacelia mollis Macbr., Soft Phacelia — MENTASTA MOUNTAINS: disturbed roadside margin, mile 34 Nabesna Road, 884 m, 62°21.18’ N 143°6.79’ W, K.A. Beck s.n., 1989; large population in rubble slope and tundra, ridge S of Soda Creek, 1173 m, 62°32.36' N 142°53.98' W, C. Roland 94-256, 22 July 1994; Dryas-forb tundra & limestone scree, Lost Creek, 1646 m, 62°34.58’ N 143°5.58’ W, M. Cook 94337, C. Roland 94-269, 23 July 1994; scattered in tundra, Boyden Hills, 1442 m, 62°30.97’ N 143°2.98’ W, C. Roland 95-088, 27 June 1995; occasional in green- stone rubble, Soda Lake, 1494 m, 62°31.2’ N 142°54.03' W, C. Roland 95-141, 5 July 1995; Dryas tundra, headwaters of Lost Creek, 1722 m, 62°36.45’ N 143°12.03’ W, K.A. Beck 95-223, 7 July 1995; scattered in dry tundra, Totschunda Creek, 1280 m, 62°27.63' N 142°12.44’ W, C. Roland 96-271, 24 June 1996; occasional in ericaceous tundra, Devils’s Mountain, 1530 m, 62°25.62’ N 142°53.93’ W, Cook & Shea 96278, 7 July 1996 (ALA); scattered in Arctostaphylos uva-ursi-Betula scrub, Lost Creek bluff, 1097 m, 62°33.3'’ N 143°9’ W, Roland & D’Auria 97-040, 26 June 1997; occasional in rocky tundra, Lost Creek, 1189 m, 62°35’ N 143°9.1’ W, Roland & D'Auria 97-060, 26 June 1997; occasional in rocky Dryas-heath-forb tundra, Lost Creek, 1646 m, 62°35’ N 143°9.1' W, Roland & D’Auria s.n., 26 June 1997; patchy in low scrub, ridge between Platinum and Totschunda Creeks, 1494 m, 62°27.89' N 142°47.38'’ W, Cook & Batten 3163, 9 July 1998 (ALA). NutzoTIn Mountains: N-facing graminoid forb tundra, 2.4 km W of Ptarmigan Creek, 1494 m, 61°55.55'’ N 141°6.72’ W, M. Cook 92465, 9 July 1992; N-facing graminoid forb tundra, 2.4km W of Ptarmigan Creek, 1494 m, 61°55.55’ N 141°6.72' W, M. Duffy 92180, 9 July 1992; rare on S- facing scree slope, Wiki Basin, 1433 m, 61°55.22’ N 141°2.75' W, M. Cook 94142, 23 June 1994; Dryas tundra slopes, headwaters of Alder Creek, 1554 m, 62°28.44’ N 142°15.06’ W, K.A. Beck 95-205, 95212, M. Cook 95132, 29 June 1995; few in bare mineral soil, Wiki Creek, 1411 m, 61°54.46’ N 141°10.68’ W, M. Cook 96100, C. Roland 96-072, 11 June 1996; bluff E of lodge, Ptarmigan Lake, 1079 m, 61°53’ N 141°8.8’ W, Duffy & Barnes 96-253, 10 August 1996; few in ericaceous heath, Wiki Creek, 1615 m, 61°55.11' N 141°11.81' W, M. Cook 3176, 15 July 1998 (ALA). St. ELIAS MOUNTAINS: occasional in ash deposit, Cub Creek, 1280 m, 61°37.27 141°9.75' W, M. Cook 94186, 28 June 1994; scat- tered over ash deposit, Mt. Natazhat, 1716 m, 61°35.38’ N 141°1.83’ W, M. Cook 95077, 20 June 41/995, This Alaska-Yukon endemic with a cordilleran distribu- tion is rare in Alaska (G2 S2S3) and in the Yukon Territory (Douglas et al. 1981). It has a wide ecological tolerance being found on disturbed roadside margins and within var- ied sub-alpine and alpine communities from mesic tundra 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 291 to steppe bluffs. The collections cited above extend its range 129 km south into the St. Elias Mountains (with intermediate stations in the Wrangell and Mentasta Mountains) from a station at Mile 1257 on the Alaska Highway (62°58.0’ N 141°38.0’ W, M. Williams 2137, 17 June 1968 (ALA)). Map 189. BORAGINACEAE Cryptantha shackletteana L.C. Higgins (C. spiculif- era (Piper) Payson), Shacklett’s Catseye — MENTASTA MOUNTAINS: scattered in red calcareous gravel on southeast to SW-facing slopes, Totschunda Creek, 1280 m, 62°27.63' N 142°12.44’ W, C. Roland 96-284, 96-295, 24 June 1996 (ALA); barren calcareous gravels slopes above Totschunda Creek, 1494 m, 62°28.04’ N 142°40.07’ W, C. Roland 96- 369, 4 July 1996 (ALA). This Alaska endemic is known from only three localities worldwide, is rare in Alaska (G1Q S1) and is a United States Fish and Wildlife Species of Concern. We have pre- viously reported this notable find (Roland and Cook 1998). This species is closely related to C. spiculifera (Piper) Payson which is common throughout the Great Basin states (Higgins 1969, Cronquist et al. 1984). A specimen from Chuktoka, assigned to C. spiculifera by Tolmachev and Yurtsev (1980), has been examined and determined to be neither C. shackletteana or C. spiculifera. The collections from the Mentasta Mountains are 280 km south of the col- lections at Eagle and Calico Bluffs on the Yukon River. Map 190. Hackelia deflexa (Wahl.) Opiz, Nodding Stickseed — St. ELtAs MOunTAINS: edge of closed tall willow scrub, Chitina River, 351 m, 6199.48’ N 142°43.57’ W, M. Duffy 91034, 3 July 1991. WRANGELL MOUNTAINS: occasional in bare soil on dry hillside, Nabesna River bluff, 0.6 km S of Bond Creek, 1106 m, 62°15.05' N 142°54.68’ W, C. Roland 96-155, M. Cook 96190, 19 June 1996. This North American boreal montane species is known from a few widely disjunct localities in Alaska and the Yukon and may be introduced (Hultén 1986; Cody 1996). The new localities cited above are from rivers some dis- tance from road systems or towns. These localities are 248- 253 km east of a station on the Glenn Highway (Hultén 1968, Welsh 1974) and 745 km west of a station in the southern Yukon Territory (Cody 1996; Welsh 1974). Map 191. SCROPHULARIACEAE Castilleja chrymactis Pennell, Green Indian Paintbrush — GULF OF ALASKA: occasional in beach strand forb herbaceous vegetation, Sudden Stream, Malaspina Forelands, 30 m, 59°47.08’ N 139°58.8’ W, M. Cook 8858, 20 July 1988; occasional in mesic tall forb meadow, Amphitheater Knob, 10 km east of Disenchantment Bay, Malaspina Forelands, 503 m, 59°57.1' N 139°46.4' W, M. Cook 87-95, 19 August 1987. This Alaska endemic with a Pacific Coastal distribution is of limited distribution in Alaska occurring from the Point Gustavus area northward to the Malaspina Forelands. This species is a consolidated hybrid between C. miniata Doug. ex Hook. and C. unalaschensis (Cham. & Schlecht.) Malte (personal communication with Mark Egger, 3 June, 2000). The stations cited above from the Malaspina Forelands are in the vicinity of collections from Disenchantment Bay (F. Funston 82, 1892 (US) (Hultén 1949)) and Yakutat Bay (Coville & Kearney 1147, 1899 (US) (Hultén 1949)). Map 192. Castilleja elegans Malte, Elegant Indian Paintbrush — CHUGACH MOUNTAINS: scattered throughout eri- caceous heath, plateau NW of Nerelna Creek, 1372 m, 61°26.66’ N 144°17.69' W, M. Cook 96297, 7 July 1996. WRANGELL MOUNTAINS: sedge tundra, Cheshnina Plateau, 500 m, 61°50.33’ N 144°22.50’ W, M. Protti 9105, M. Duffy 91086, 21 July 1991; mesic herbaceous/dwarf scrub tundra, ridge between Chichokna and Chetasline Rivers, 1463 m, 61° 56.51’ N 144° 25.93’ W, M. Cook 94471, 15 August 1994; top of small ridgetop, Chokosna River, 1097 m, 61°32.46’ N 143°32.88' W, J. Bolivar 84-101, 13 July 1994; Dryas sedge tundra, NW slope of Mt. Sanford between Sanford and Boulder Creeks, 395 m, 62°21.17’ N 144°24.52’ W, M. Cook 93122, 9 June 1993; scattered in Dryas-graminoid tundra, NW slopes of Mt. Sanford, 1387 m, 62°20.98’ N 144°28.18' W, M. Cook 94411, 30 July 1994; occa- sional in forb herbaceous meadow, vic. Nadina Glacier, 1768 m, 62°2.85’ N 144°41' W, M. Cook 94422, 30 August 1994; common in heath snowbed tundra, volcanic ridge 6 km east of Snyder Peak, 1524 m, 62°4.47' N 144°30.51' W, C. Roland 96- 377, 5 July 1996; scattered in sparse, lichen-domi- nated heath tundra, Cheshnina Plateau, 1399 m, 61°48.04' N 144°6.27' W, C. Roland 96-871, 96- 877, 5 August 1996; Cassiope-Dryas heath, plateau between Long Glacier and the Cheshnina River, 1399 m, 61°48.04’ N 144°6.27' W, M. Cook 96718, 8 August 1996. This is an amphiberingean species with an arctic-alpine distribution. The collections cited above extend its range 312 km south into the Chugach Mountains and 244 km south into the Wrangell Mountains from a collection in the Alaska Range (Mt. Hayes Quad: 63°40.0’ N 144°54.0' W, L.A. Spetzman 86, 20 June 1957 (ALA)). There may have been two collections in the Wrangell Mountains (Jacksina River, Schrader & Hartman 49, 1902 (US) and Sanford River, W.L. Poto 100, 1902 (US) (Hultén 1948)). Hultén determined these specimens as C. pallida (L.) Kunth subsp. mexiae Pennell in the Flora of Alaska and the Yukon (1949). Hultén followed the treatments of B. Boivin (1952) and O.V. Rebristaja (1964) in his 1968 flora in which C. elegans is regarded as a distinct species and C. pallida ssp. mexiae is united with C. elegans (Hultén 1967, 1968). However, the distribution map in the 1968 flora for C. ele- gans does not include the specimens from the Wrangell Mountains or other specimens of C. pallida ssp. mexiae as cited in the Flora of Alaska and the Yukon (1949) nor do they appear to be included on the map of C. caudata (Pennell) Rebr. If these stations were included in the current distribution of C. elegans, it would extend south to Keystone Canyon on the Richardson Highway (J.P. Anderson 1884 (S)), west to Matanuska (J.P. Anderson A TT 202 1908 (NY)) and include a locality near Willow Creek along the Richardson Highway adjacent to the Park (J.P. Anderson 2003 (S) (Hultén 1949)). Map 193. Castilleja yukonis Pennell, Yukon Indian Paintbrush — WRANGELL MOUNTAINS: scattered on rocky W- facing bluff in open alder scrub, between Nabesna River and Ellis Lake, 1310 m, 62°16.22' 142° 55.70’, M. Cook 3026, 21 June 1997. This species is an Alaska- Yukon endemic with a boreal- montane distribution. The specimen cited above represents the western extent of its distribution and extends its range 90 km into the northwest Wrangell Mountains from a collection at Skolai Pass (61°37.25'’N 141°58.77'W, D. F. Murray 1025 (CAN) (Murray 1968)). The closest station to the Wrangell Mountain collections is 190 km to the east in the Yukon Territory (Cody 1996). The collections from the Wrangell Mountains are the only two known localities from Alaska. Map 194. Euphrasia mollis (Ledeb.) Wettst., Subalpine Eyebright — ST. ELIAS MOUNTAINS: White spruce forest, Clear Stream, 61°6.31'N 141°58.88' W, M. Hoffman s.n., 11 August 1986. CHUGACH MOUNTAINS: sand, Bremner River, 1300 m, 60°54.71' N 144°33.33’ W, M. Duffy 91138, 16 August 1991. This species is amphiberingean with a North Pacific coastal distribution. The collections cited above extend its range 205 km east into the St. Elias Mountains from collec- tions near Cordova (Alaganik Slough 60°24.16'’N 145°28.45'W, E. Helmstetter s.n., 8 August 1980 (ALA)) and are 186 km northwest of collections at Yakutat (Situk, 59°40.59'N 139°37.38'W, M.C. Muller s.n., 17 July 1980 (ALA)). Map 195. Penstemon gormanii Greene, Gorman’s Beardtongue — WRANGELL MOUNTAINS: small population in lime- stone rubble at base of hill 3070’, 1.2 km SE of Whitham Lake, 936 m, 62°19.33’ N 142°53.28’ W, C. Roland 96-177, M. Cook 96198, 20 June 1996; abundant on small knob undercut by the Nabesna River downstream of Virginia lake, 759 m, 62°21.51' N 142°52.87' W, C. Roland 96-215, M. Cook 96230, 21 June 1996; abundant in bare soil on steep, S-facing slopes, SW end of Devil’s Mountain, 942 m, 62°24.95’ N 142°54.86' W, C. Roland 96- 227, M. Cook 96234, 22 June 1996; scattered in moss on limestone ledges and in crevices, Nabesna River at Cheslina River, 677 m, 62°47.49' N 142°10.4’ W, M. Cook 96260, 25 June 1996. This is an Alaska-Yukon endemic with a boreal-mon- tane distribution. The collections cited above extend its range 80 km south into the Wrangell Mountains from a sta- tion on the Slana-Tok Highway in the Mentasta Mountains (62°36.0’ N 144°38.0' W, J.P. Anderson 8692, 23 June 1944 (ALA)). Map 196. Synthyris borealis Pennell, Northern Kittentails — MENTASTA MOUNTAINS: Upper Trail Creek, 1341 m, 62°37.09' N 143°16.06' W, Leggett & Potkin 95- 045, 26 July 1995; occasional in fine gravel on E- facing slope, Totschunda Creek, 1280 m, THE CANADIAN FIELD-NATURALIST Vol. 116 62°27.63' N 142°12.44’ W, C. Roland 96-294, 24 June 1996. NUTZOTIN MOUNTAINS: few in volcanic boulder field, Wiki Peak, 1411 m, 61°54.46’ N 141°10.68’ W, M. Cook 96096, 10 June 1996; S-fac- ing scree, Ptarmigan Creek, 1494 m, 61°32.89’ N 141°3.28' W, M. Cook 92476, 9 July 1992; talus slope, Gold Hill Peak 5745’, 1585 m, 62°6.73’ N 141°51.82' W, M. Cook 9084, 24 July 1990; seral herbs on well vegetated SE-facing alpine colluvium, 1128 m, 62°2’ N 141°11' W, Parker & Gracz 6912, 13 August 1996; abundant on N-facing talus slope, 1411 m, 61°54.46’ N 141°10.68’ W, C. Roland 96- 077, 7 June 1996; moist alpine herbs on NW-facing slope, vic. Wiki Peak, 1585 m; 61°33,98 3) 141°8.02' W, C. Roland 94-118A, 23 June 1994. WRANGELL MOUNTAINS: lower slopes of cinder cone, Cone Ridge, 2073 m, 62°8.21’ N 143°18.47’ W, M. Cook 94409, C. Roland 94-320, 27 July 1994; scat- tered in unstable basalt scree, Black Mountain, 1481 m, 62°20.85’ N 143°44.9’ W, M. Cook 95148, 2 iuky 1993: This species is an Alaska-Yukon endemic with a cordilleran distribution. It had been collected in the St. Elias Mountains at Guerin Glacier (61°37.42’N 141°4.38’W, D.F. Murray 2020, 3 August 1968 (ALA)). The new collections extend its range northwest 106 km north into the Nutzotin Mountains, 150 km into the Mentata Mountains and 153 km west into the Wrangell Mountains. These localities connect its distribution 195 km to the northwest in the Alaska Range (Hultén 1968) with the dis- tribution to the east in the Yukon Territory (Cody 1996). Map 197. Veronica serpyllifolia L. ssp. humifusa (Dickson) Syme, Thyme-leaf Speedwell — Sr. ELIAS MOUNTAINS: open tall willow scrub, Guyot Glacier, 60°5.43’ N 141°25.51’ W, M. Duffy 92254, 22 August 1992; dry forb alpine herbaceous alluvial slope, Samovar Hills, 518 m, 60°8.35’ N 140°39.6' W, M. Cook 87-151, 23 August 1987. This species is incompletely circumpolar with a widespread distribution. It had been collected in the vicini- ty of the Park in Disenchantment Bay (Coville & Kearney s.n., June 1899 (US) (Hultén 1947)). The collections cited above connect the coastal range 313 km to the west with the collection in Disenchantment Bay 104 km to the east. Map 198. PLANTAGINACEAE Plantago eriopoda Torr., Saline Plantain — COPPER RIVER BASIN: sandy riverbed, Copper River, 381 m, 62°10.5' N 145°24.57' W, K. Teare 1659, 8 June 1984; alluvium near margin of lower Klawasi mud volcano, 563 m, 62°3.52’ N 145°13.29’ W, M. Cook 91216, 21 August 1991; mud volcano salt pan, 62°3.69’ N 145°12.49’ W, M. Protti 9101, 6 June .1991; meadow, Lower Klawasi River, 366 m, 62°1.8’ N 145°20.8’ W, C. Roland 94-001, 29 May 1994. This North American species is disjunct from the tem- perate zone in the southwest Yukon and adjacent southcen- tral Alaska. The stations listed above are 200 km to the 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 293 southwest of a station near Northway (Hultén 1968) and represent the western extent of its distribution. Map 199. ASTERACEAE Agoseris glauca (Pursh) Raf., Pale Agoseris — CHUGACH MOUNTAINS: mesic forb tundra, Upper Tebay Lake, 1219 m, 61°12.18’ N 144°23.8’ W, M. Duffy 91113, 12 August 1991; meadow near the Bremner River, 396 m, 60°58.98’ N 144°17.1' W, . Duffy 91146, 19 August 1991. This North American boreal-montane species is rare in Alaska (G4G5 S1) where it reaches the western extent of its range. The specimens cited above from the Chugach Mountains connect its distribution with one locality 391 km to the southwest (Seldovia Quad: 59°40.0’ N 151°09.0' W, E. Berg 47, 11 August 1988 (ALA)) with collections 478 km to the southeast near Skagway (Hultén 1968). Map 200. Antennaria media Greene, Rocky Mountain Pussytoes — CHUGACH MOUNTAINS: moss-forb herbaceous slope, alpine valley near Hanagita Peak, 1186 m, 61°4.91’ N 143°38.86' W, M. Cook 96415A, 7 July 1996; occasional in Cassiope-Luetkea heath of alpine basin, vic. 12-mile Creek, 1271 m, 60°48.85’ N 142°33.52’ W, M. Cook 96625, 7 July 1996 (ALA); rare in sand deposits near shore of Rock Lake, 1119 m, 61°48.7’ N 141°16.57’ W, C. Roland 96-056, 7 June 1996. ST. ELIAS MOUNTAINS: occasional in tundra gravel patches, headwaters of Chitina River between Walsh and Logan Glaciers, 1951 m, 60°53.94’ N 141°6.75’ W, C. Roland 95- 166, 12 July 1995. This species is North American with a cordilleran distri- bution. It is rare in Alaska and in the Yukon (Bayer 1993). The specimens cited above are 95 km northeast of the the only other Alaskan locality at Cordova (60°58.0’ N 145°00.0' W, C. Parker 1865, 12 August 1986 (ALA)) and are 82 km east of collections in the Yukon Territory (Cody 1996). Map 201. Arnica amplexicaulis Nutt. ssp. prima Maguire, Streambank Leopardbane — CHUGACH MOUNTAINS: mesic forb tundra, upper Tebay Lake, 1219 m, 61°12.18’ N 144°23.8' W, M. Duffy 91107, 12 August 1991. GULF OF ALASKA: closed tall scrub, glacial barren, vic. Tann Fjord, Icy Bay, 305 m, 60°7.52' N 141°8.1' W, B. Haller s.n., 24 August 1987. This is a North American cordilleran species considered rare in the Yukon Territory (Douglas et al. 1981). The localities cited above connect the range 115 km to the southwest near Cordova (Hultén 1968) with the range 197 km to the east in the Yukon Territory (Cody 1996) and 110 km to the southeast at Yakutat (Hultén 1968). Map 202. Arnica latifolia Bong., Broadleaf Arnica — CHUGACH MOUNTAINS: steep, N-facing snowflush slope, Granite Creek, 579 m, 60°44’ N 142°32' W, Parker & Duffy 6712, 6 August 1996; open low willow scrub, Blue Lake, 936 m, 61°17.94' N 144°10.71' W, M. Cook 96364, 7 July 1996; patchy in mesic forb meadow, 12-mile Creek, 1326 m, 60°50.21’ N 142°30.85’ W, M. Cook 96596, 7 July 1996; scattered in mixed forb herbaceous meadow, Granite River, 1344 m, 60°46.44' N 142°5.12' W, M. Cook 96619, 7 July 1996; Empetrum nigrum/mixed forb community on wind-exposed rocky area, Falls Creek, 655 m, 61°14.17' N 144°28.24' W, L. A. & E. G. Viereck 11028, 7 July 1996; scattered in subarctic lowland sedge wet meadow, Tebay Lake, 579 m, 61°11’ N 144°24’ W, Parker & Gracz 6745, 7 August 1996; Betula glandulosa thicket, Martin Creek, 1097 m, 60°56’ N 142°23' W, Batten & Barker 96-256, 28 July 1996; lush graminoid-forb meadow, Spirit Mountain, 759 m, 61°20.76’ N 144°26.89’ W, C. Roland 96-564, 15 July 1996. WRANGELL MOUNTAINS: common in lush streamside meadow, Mill Creek, 1241 m, 61°33.33’ N 143°28.35’ W, C. Roland 96-661, 24 July 1996; locally abundant in lush meadow, Castle Peak, 1219 m, 61°33.64’ N 143°19.01’ W, C. Roland 96-692, 96-701B, 25 July 1996; clumped in mesic tall forb-graminoid herba- ceous meadow, Nadina Glacier, 1768 m, 62°2.85’ N 144°41’ W, M. Cook 94417, 30 August 1994. Sr. ELIAS MOUNTAINS: herbaceous meadow, Karr Hills, 549 m, 60°8.66’ N 141°16.44’ W, K. Beck s.n., 12 August 1987. This Arnica is North American with a cordilleran distri- bution. The collections cited above extend its range 141 km east into the Wrangell Mountains and 244 km southeast into the Chugach Mountains from a station on the Glenn Highway near Glennallen (Hultén 1968). The southern St. Elias Mountain locality connects the range 241 km to the west near Cordova (Hultén 1968) with a station 121 km to the east at Yakutat (Hultén 1968). Map 203. Arnica mollis Hook., Hairy Arnica — WRANGELL MOUNTAINS: forb herbaceous meadow, West Fork Mill Creek, 1241 m, 61°33.33’ N 143°28.35’ W, C. Roland 96-659, 24 July 1996. This is a North America cordilleran species which is rare in Alaska (G5 S1) and in the Yukon Terriotry (Douglas et al. 1981). The specimen cited above extends the range into the Wrangell Mountains and connects the distribution of this species 265 km to the west in the Anchorage Quad (Hatcher Pass, 61°46.0’ N 149°18.0’ W, T. Ward 95, 14 August 1976 (ALA)) with the distribution 445 km to the east in the Yukon Territory (Cody 1996). Map 204. Artemisia hyperborea Rydb. (A. furcata auct. non Bieb.), Three-Forked Wormwood, — MENTASTA MOUNTAINS: occasional on S-facing slope in lime- stone gravels near ridge, Soda Lake, 1173 m, 62°32.36' N 142°53.98' W, C. Roland 94-259C, 22 July 1994; occasional on limestone outcrop between Soda and Platnium Creeks, 1494 m, 62°31.2' N 142°54.03' W, C. Roland 95-139, 5 July 1995; scat- tered in rocky tundra and steppe on S-facing slopes, Devil's Mountain, 942 m, 62°24.95' N 142°54.86' W, C. Roland 96-241, 22 June 1996; common on dry slopes, Totschunda Creek, 1280 m, 62°27.63' N 142°12.44' W, C. Roland 96-273, 24 June 1996; —— ee ewe eS eS - ” 294 THE CANADIAN FIELD-NATURALIST Vol. 116 191. Hackelia deflexa 192. Castilleja chrymactis 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 196. Penstemon gormanii 205 ee ee eee ee Oi 296 crevices of S-facing rock outcrop on spur ridge crest between Platinum and Totschunda Creeks, 1400 m, 62°27.83' N 142°46.23’ W, A.R. Batten 98-36; south side of ridge on exposed rock outcrops, ridge between Platinum and Totschunda Creeks, 1524 m, 62°27.82'’ N 142°46.27' W, M. Cook 3157, 9 Suly 1998. NUTZOTIN MOUNTAINS: common on disturbed granitic rubble, vic. Carl and James Creek conflu- ence, 1920 m, 62°3.52' N 141°36.27' W, C. Roland 95-024, 14 June 1994; Wiki Peak ridge, 1433 m, 61°53.9’ N 141°9.51' W, M. Cook 94127, 22 June 1994; occasional in open soil, ridge 1.8 km N of Ptarmigan Lake, 1433 m, 61°53.9’ N 141°9.51’ W, C. Roland 94-111B, 22 June 1994; abundant on SW- facing scree slope, Horsfeld Creek, 1768 m, 62°2.88' N 141°13.18’ W, C. Roland 94-131B, 24 June 1994; common on steep, S-facing rubble slope, Klein Creek, 1747 m, 62°2.29’ N 141°19.88’ W, C. Roland 95-037, 15 June 1995; volcanic plateau between Bryan and Willow Creeks, 1829 m, 61°58.97' N 141°50.03’ W, M. Cook 95053, 18 June 1995; scat- tered in dry soil of subalpine steppe, Wiki Basin, 1411 m, 61°54.46’ N 141°10.68’ W, C. Roland 96- 046B, 96-110, M. Cook 96062, 11 June 1996; patchy in rocky steppe vegetation on S-facing slopes, Rock Lake, 1119 m, 61°48.7’ N 141°16.57' W, C. Roland 96-011, 96-011, 4 June 1996; patchy in open low birch/willow shrub, Francis Creek, 1280 m, 61°53.29' N 141°13.1’ W, M. Cook 3169, 14 June 1998; few on unstable gravel and cobble S-facing rhyolite scree slope, vic. Wiki Creek, 1524 m, 61°54.77' N 141°11.05' W, M. Cook 3182, 15 July 1998; scattered in gravel sized scree on basalt dome, Sonja Creek, 1646 m, 61°57.17' N 141°21.92’ W, M. Cook 3195, 16 July 1998. WRANGELL MOUNTAINS: occasional on SW-facing talus slope, White River and Lime Creek confluence, 1646 m, 61°46.25’ N 141°50.36’ W, M. Cook 94105A, 21 June 1994; scat- tered on S-facing slope, Capitol Mountain, 1052 m, 62°31.61' N 144°12.38' W, C. Roland 94-014, 7 June 1994; locally abundant on steep, S-facing slope, Fish Creek, 1067 m, 62°16.92’ N 142°58.99' W, C. Roland 95-303, 30 July 1995; common on S-facing cutbank, Mud, Lake, 100k m, 62°13.52’ N 143°45.28’ W, C. Roland 96-894, 10 August 1996. This species is an Alaska-Yukon endemic with a cordilleran distribution. It was collected in the St. Elias Mountains (Sheep Glacier, 61°42.48'N 141°38.59'W, D. F. Murray 2232, 15 August 1968 (ALA)). The collections cited above extend the range 163 km northwest into the Wrangell Mountains, 116 km northwest into the Mentasta Mountains and 89 km northwest into the Nutzotin Mountains from the Sheep Glacier locality. These new localities connect the distribution of this species 114 km to the northwest in the Alaska Range (Gunnysack Creek, 63°32.0’ N 145°51.0' W, Batten et al. 78-28, 28 June 1978 (ALA)) with the distribution 76 km to the east in the Yukon Territory (Cody 1996). Map 205. Aster alpinus L. ssp. vierhapperi Onno, Alpine Aster — CHUGACH MOUNTAINS: forb herbaceous meadow, THE CANADIAN FIELD-NATURALIST Vol. 116 alpine basin between Twelve-mile Creek and Tana River, 1280 m, 60°49.8’ N 142°33.3' W, M. Hoffman s.n., 7 August 1986. This North American cordilleran species is known from only eight localities in Alaska, six of these localities are quite disjunct. The collection cited above extends its range 388 km south into the Chugach Mountains from a collec- tion in the Big Delta Quad (confluence of the Tanana & Delta Rivers, 64°09.0’ N 145°50.0' W, R. Howenstein s.n., 5 August 1983 (ALA)) and connects the Alaska range with the distribution in the Yukon Territory 157 km to the east where it is more widespread. Map 206. Aster borealis (T. & G.) Provancher (A. junciformis Rydb.) — WRANGELL MOUNTAINS: occasional in Myrica gale dominated quaking Drepanocladus moss mats, 1.5 mi northwest of Billy Lake, McCarthy Road, 614 m, 61°27.99’ N 143°56.38’ W, M. Cook 94453, 10 August 1994. The specimen cited above extends the range of this North American boreal-montane species 220 km west into the Wrangell Mountains from collections in the Anchorage Quad (Potter Point, 61°60.0'’ N 149°55.0’ W, Batten & Murphy 77-474, 8 August 1977 (ALA)) and connects the distribution 189 km to the east in the Yukon Territory (Cody 1996). Map 207. Erigeron caespitosus Nutt., Tufted Fleabane — MENTASTA MOUNTAINS: scattered on sparsely vege- tated slope in sandy gravel, Devil's Mountain, 942 m, 62°24.95’ N 142°54.86' W, M. Cook 96238, C. Roland 96-228, 22 June 1996 (ALA); occasional in dry rock crevices and rubble on SE-facing slope in midgrass herb vegetation, Devil’s Mt., 1530 m, 62°25.62' N 142°53.93’ W, C. Roland 96-354, M. Cook 96282, 96283, 3 July 1996; scattered on boul- der slope, 500 m above treeline, ridge between Platinum and Totschunda Creeks, 1494 m, 62°27.82’ N 142°46.27' W, M. Cook 3161, 3164, 9 July 1998. NUTZOTIN MOUNTAINS: abundant on gravel substrate in mixed forb herbaceous vegetation, Chisana airstrip, 1006 m, 62°4.16’ N 142°2.84’ W, M. Duffy 91096, 24 July 1991; common on SW-facing bluff, ridge 0.6 km E of Rocker & Ptarmigan Creek conflu- ence, 1311 m, 61°54.8’ N 141°2.03’ W, C. Roland 94-121, 23 June 1994; common on S-facing bluff between Cub and Traver Creeks, White River, 1219 m, 61°44.3’ N 141°9.5'’ W, C. Roland 95-081, 21 June 1995; midgrass-herbaceous S-facing bluffs, Ptarmigan Lake, 1128 m, 61°50.12' N 141°9.15’ W, Duffy & Barnes 96-289, 8 August 1996. WRANGELL MOUNTAINS: locally abundant on steep, S-facing slope, Fish Creek, 1067 m, 62°16.92' N 142°58.99’ W, C. Roland 95-301, 30 July 1995; common on steep SE-facing slope in sagebrush-grass vegetation, Nabesna River, 0.6 km S of Bond Creek, 1106 m, » 62°15.05' N 142°54.68' W, C. Roland 96-167, 19 June 1996. This North American species is disjunct in Alaska and the Yukon from the Rocky Mountains. It had been collect- ed on the Nabesna River (Schrader & Hartman 65, 1902 (US) (Hultén 1948)) and in Chitistone Pass (61°37' N 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 297 142°3’ W, R. Scott s.n., 1967 (MICH) (Scott 1968)). The stations cited above extend its range 64 km into the Nutzotin Mountains and connect its distribution 69 km to the north along the Alaska Highway in the Tanana Valley (Hultén 1968) with the distribution 111 km to the east in the Yukon Territory (Cody 1996). Map 208. Erigeron grandiflorus Hook. ssp. arcticus A.E. Porsild (E. grandiflorus Hook. ssp. grandiflorus), Large-Flowered Fleabane — CHUGACH MOUNTAINS: rare in meadow, plateau above Nerelna Creek, 1372 m, 61°26.66’ N 144°17.69' W, Roland & D’Auria 96-420, 8 July 1996. MENTASTA MOUNTAINS: E-fac- ing ravine at crest on limestone ridge south of Soda Lake, 1173 m, 62°32.36' N 142°53.98’ W, M. Cook 94305, 21 July 1994; scattered on SW-facing out- crop near ridge in mesic graminoid forb herbaceous vegetation, Soda Lake, 1173 m, 62°32.36’ N 142°53.98’ W, M. Cook 94325, C. Roland 94-2594, 22 July 1994; scattered on outcropping, south side of limestone ridge, Lost Creeks, 1646 m, 62°34.58’ N 143°5.58' W, M. Cook 94328, 23 July 1994; scat- tered in sheep beds on S-facing outcrop in graminoid-forb-dwarf scrub vegetation, headwaters of Lost Creek, 1722 m, 62°36.45’ N 143°12.03’ W, M. Cook 95182, 7 July 1995; confined to dry forb- graminoid-Dryas tundra slopes above mouth of Totschunda Creek, 1280 m, 62°27.63’ N 142°12.44’ W, C. Roland 96-274, 24 June 1996; scattered in turfy graminoid tundra amid cobble-sized clastic talus, E-facing slope, Devil's Mountain, 1530 m, 62°25.62' N 142°53.93’ W, C. Roland 96-349, 3 July 1996. NUTZOTIN MOUNTAINS: dry sandy soil on ridge, Carl Creek, 1920 m, 62°3.52' N 141°36.27’ W, M. Cook 95007, 14 June 1994; steep, disturbed and xeric gravel slope, ridge 4 km north of Ptarmigan Lake, 1433 m, 61°53.9' N 141°9.51' W, C. Roland 94-117A, 22 June 1994; occasional on SW-facing slope, headwaters of Alder Creek, 1554 m, 62°28.44' N 142°15.06' W, K.A. Beck 95- 213, 30 June 1995. St. ELIAS MOUNTAINS: rare in SW-facing limestone gravels above cliffs, ridge between Dan & Copper Creeks, 1554 m, 61°21.56' N 142°26.43' W, C. Roland 94-240, 14 July 1994. WRANGELL MOUNTAINS: rare on turfy, sheep grazed site in organic rich soil of Dryas tundra, Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85’ W, C. Roland 95-239, 18 July 1995; N-facing scree slope, Lime Creek, 1707 m, 61°28.41’ N 141°29.44' W, M. Cook 92496, 7 July 1992; SE-facing shaley scree near ridge, Nikolai Mine, 1695 m, 61°27’ N 142°39' W, Batten & Barker 96-066, 24 July 1996. This species is an Alaska-Yukon endemic with an arc- tic-alpine distribution. It had been collected in the St. Elias Mountains (Guerin Glacier, 61°37.42'N 141°4.38'W, D.F. Murray 2080, 4 August 1969 (ALA)). The stations cited above extend its range from the St. Elias Mountains 157 km to the northwest into the Mentasta Mountains, 113 km northwest into the Nutzotin Mountains, 133 km southwest into the southern Wrangell Mountains and 170 km south- west into the Chugach Mountains. These new localities connect its distribution 185 km to the west in the Alaska Range (Hultén 1968) with the distribution 48 km to the east in the Yukon Territory (Cody 1996). Map 209. Saussurea angustifolia (Willd.) DC. ssp. yukonensis (Porsild) Cody (S. viscida Hult. var. yukonensis (Pors.) Hult.), Narrowleaved Saussurea — CHUGACH MOUNTAINS: Cassiope tundra, Towhead Mountain, 1433 m, 61°3.21’ N 142°39.8' W, Duffy & Barnes 96-149, 7 August 1996; rock outcrops, Nelson Mountain, 1558 m, 61°19.37' N 143°48.83’ W, M. Cook 96378, 7 July 1996. MENTASTA MOUNTAINS: Trail Creek, 1341 m, 62°37.09’ N 143°16.06’ W, Potkin & Leggett 95-046, 27 July 1995; limestone scree, ridge between Lost and Platinum Creeks, 1646 m, 62°34.58’ N 143°5.58' W, M. Cook 94340, 23 July 1994. WRANGELL MOUNTAINS: Lakes Plateau, 1890 m, 62°4.4' N 143°23.5' W, Potkin & Leggett 95-114, 30 July 1995; rubble slope, Cone Ridge, 2073 m, 62°8.21' N 143°18.47' W, M. Cook 94406, 26 July 1994; scree slope, Lime Creek, 1707 m, 61°28.41’ N 141°29.44’ W, M. Duffy 92209, 11 July 1992; common in clay soil on E-facing slope, Grotto Creek, 1847 m, 61°30.56"°.N 142°24.79" W, C. Roland 96-648, 23 July 1996; bare gravel chert, Lime Creek, 1646 m, 61°46.25’ N 141°50.36’ W, C. Roland 94-105, 21 June 1994. This species is an Alaska-Yukon endemic with an arc- tic-alpine distribution. It had been collected in the Wrangell-St. Elias Mountains at Bonanza Ridge (Nordell and Schmitt 1978), Sheep Glacier (D.F. Murray 2239 (ALA) 1968), Guerin Glacier (D.F. Murray 2089, 2027, (ALA) 1968) and Chitistone Pass (Scott 1968). The collec- tions cited above extend its range 134 km northwest into the Mentasta Mountains and 139 km west into the western Wrangells and connect its distribution 207 km to the north- west in the Alaska Range (Hultén 1968) with its range in the Yukon Territory (Cody 1996). Map 210. Taraxacum carneocoloratum Nels., Pink Dandelion — MENTASTA MOUNTAINS: rare on N-facing scree slope in Nikolai Greenstone and limestone units, Soda Lake, 1173 m, 62°32.36’ N 142°53.98' W, M. Cook 94301, 21 July 1994; few in reddish gravel scree, SE-facing slope above Totschunda Creek, 1494 m, 62°28.28' N 142°40.5’ W, K. Shea 96287, 7 July 1996; ridge above Totschunda Creek, few 5 m below ridge in sparsely vegetated orange gravel scree, 1451 m, 62°29.02’ N 142°43.94' W, M. Cook 96293, 7 July 1996. NUTZOTIN MOUNTAINS: ridge 1.6 km W of Ptarmigan & Rocker Creek confluence, alpine tundra and rock slope, 1615 m, 61°54.81' N 141°5.51' W, M. Duffy 92187, 9 July 1992; ridge at headwaters of Alder Creek, occasional on S-facing boulder and rubble slopes, 1554 m, 62°28.44' N 142°15.06' W, K.A. Beck 95-216, 29 June 1995. ST. ELIAS MOUNTAINS: Mt. Chitina, rare in steep, loose scree at contact between limestone and darker meta- morphic rock, 2073 m, 60°57.74' N 141°17.33' W, C. Roland 95-224, 16 July 1995. WRANGELL MOUNTAINS: Lime Creek, alpine scree slope, 298 THE CANADIAN FIELD-NATURALIST Vol. 116 200. Agoseris glauca 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 201. Antennaria media 204. Arnica mollis 209 eee eee eee eee —™ —— - O 300 THE CANADIAN FIELD-NATURALIST Vol. 116 208. Erigeron caespitosus 2002 COOK AND ROLAND: NOTABLE VASCULAR PLANTS FROM ALASKA 211. Taraxacum carneocoloratum 212. Taraxacum phymatocarpum 301 —--- SO —-—™”™ —_— SO 302 1935 m, 61°47.35' N 141°49.06' W, Duffy & Cook 92106, 8 August 1992; Cooper Pass, rare in moist orange colored clay soil between gravel and thin bedded talus shale on N-facing slope, 1942 m, 62°17.16’ N 142°31.44' W, M. Cook 94357, 24 July 1994; rare in rocks and gravel on ridgeline, Crystalline Hills, 1585 m, 61°23.59’ N 143°31.85’ W, C. Roland 95-242, 18 July 1995; mesic rubble slope, Jaegar Mesa, 1893 m, 62°15.9’ N 143°1.24' W, Moran & Roland 95-45, 8 August 1995. This Alaska- Yukon endemic with a cordilleran distribu- tion is known from one locality in the Yukon Territory and sixteen localities in Alaska, ten of which are cited above from the Park. These collections are 273 km east of a sta- tion in the Anchorage Quad (Campbell Lake, 61°07.17' N 149°29.75’ W, Lichvar et al. 8082A, 19 July 1994 (ALA)) and 409 km south of the station in the Ogilive Mountains, Yukon Territory (Murray and Lipkin 1987). Hultén (1973) indicated that this species may be the same as T. soczavae Tzvelev, whereas Welsh (1974) considered it to be a vari- ant of T. eriophorum Rydberg (Murray and Lipkin 1987). Map 211. Taraxacum phymatocarpum J. Vahl (Taraxacum lyratum (Ledb.) DC. in part), Northern Dandelion — CHUGACH MOUNTAINS: scree, West Fork Goat Creek, 1487 m, 60°59.5’ N 142°1’ W, Batten & Barker 96- 301, 29 July 1996 (ALA). This species is amphiberingean with an arctic-alpine dis- tribution. The collection cited above extends the range 554 km south into the Chugach Mountains from a station in the Steese-White Moutains (Hultén 1968) and connects the range 164 km to the east in the Yukon Territory (Cody 1996). Map 212. Acknowledgments This inventory was funded by the U.S. National Park Service Natural Resource Preservation Program. Mike Duffy was essential in developing the proposal for this inventory through his extensive collecting and knowledge of the flora of the region. For conducting field inventories we thank: Allan Batten, Marilyn Barker, Jennifer Barnes, Katy Beck, Mike Duffy, Mike Gracz, Anne Leggett, Virginia Moran, Carolyn Parker, Michele Potkin, Leslie Viereck and Eleanor Viereck. National Park Service staff who facilitated this study include: Vicki Ables, Geoff Bleakley, Russell Galipeau, Sue Huse, Devi Sharp and Anne Worthington. We are grateful to Carolyn Parker for reviewing most of our specimens, Allan Batten for providing collection data from the University of Alaska Northern Plant Documentation Center and David F. Murray for reviewing difficult taxa and providing advice throughout this inventory. We would also like to acknowledge the following specialists for examining our specimens and provid- ing determinations: George Argus, Mark Egger, Reidar Elven, Barbara Ertter, Donald Farrar, Signe Fredericksen, G.A. Mulligan, Robert Soreng and Marcia Waterway. 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A collection of plants from Yakutat, Alaska. Bartonia 24: 9-21. U.S. Department of the Interior. 1986. General manage- ment plans for Wrangell-St. Elias National Park and Preserve, Alaska. 239 pages. U.S. Department of the Interior. 1991. NPS-77, Natural resources management guidelines. Welsh, S. 1974. Anderson's flora of Alaska and adjacent parts of Canada. Brigham Young University Press, Provo, Utah. 724 pages. Yurtsev, B. A. 1963. On the floristic relations between steppes and praires. Botaniska Notiser 116: 396-408. Received 19 April 2001 Accepted 4 July 2002 Notes Black Bear, Ursus americanus, Hair and Apple Trees, Malus pumila, in Northeast North America DAVID H. HirTHuH!, Jos—epH M. A. PETry!, and C. WILLIAM KILPATRICK2 \Wildlife and Fisheries Biology Program, University of Vermont, Burlington, Vermont 05405 USA 2Department of Biology, University of Vermont, Burlington, Vermont 05405 USA Hirth, David H., Joseph M. A. Petty, and C. William Kilpatrick. 2002. Black Bear, Ursus americanus, hair and Apple Trees, Malus pumila, in northeast North America. Canadian Field-Naturalist 116(2): 305-307. Molecular genetic techniques make it possible to use extremely small DNA samples for genetic analysis of Black Bears (Ursus americanus). Non-invasive sources of DNA include feces and hair. Compared to feces, hair has the advantage of being a stable source of DNA and may be taken from living or dead bears. A number of researchers have collected hair in “hair traps”, where bears are induced to reach across barbed wire to get at a bait. However, we have discovered that in the Northeast bears climb wild Crab Apple trees (Malus pumila) in the fall for fruit and leave hairs on bark and broken twigs in the process. In this part of the United States and Canada, feral Apple trees appear to be a productive source of bear hair that has application for genetic studies. Key Words: Black Bear, Ursus americanus, genetics, Wild Crab Apple tree, Malus pumila. The combination of modern molecular genetics and non-invasive collection of genetic material represents one of the most exciting advances in field biology since the development of radiotelemetry in the 1960s and 1970s (Parker et al. 1998; Taberlet and Waits 1998; Woods et al. 1999). Using microsatellite tech- niques and DNA samples obtained from tissue, feces, or hair, geneticists are able to identify individuals, sex, members of immediate family groups, members of more distant breeding populations, as well as estimate size of effective breeding population (Taberlet et al. 1993; Paetkau and Strobeck 1994; Haig 1998; Waples 1989). A key aspect in these procedures has been the development of the polymerase chain reaction (PCR), which makes it possible to work with very small genetic samples (Taberlet et al. 1996). Information obtained from these genetic analyses may be particularly important in field studies of ani- mals that: (1) have large home ranges, (2) occur at low densities, or (3) are difficult to capture and mark. Large home ranges (>250 km?) make conven- tional radiotelemetry difficult because animals often move beyond the range of receivers. Animals that occur at low densities may be difficult to locate or attract and are therefore problematic for capture and radiotelemetry. Capture of some large mammals, such as bears or large cats, may involve inherent risks to researchers or study animals through use of drugs or dogs. Black Bears (Ursus americanus) exemplify all of the above difficulties. They occur at low densities (e.g., 1 bear per 6 km? in Minnesota [Rogers 1987], 1 bear per 30 km? in Michigan [Whitaker and Hamilton 1998]). They are shy, solitary, and almost impossible to observe in the field. Young male bears disperse long distances from their mother’s home range (>100 km) (Pelton 2000), and male bears in particular have large home ranges (e.g., 318 km? in Massachusetts [Elowe 1984], 170 km2 in Pennsy]l- vania [Alt et al. 1980]). Once attracted to bait or treed by dogs, the bear must be drugged to be han- dled, which carries an inherent risk to the bear. One advantage of genetic techniques is that DNA samples can be taken from a number of sources and incorporated into the same analysis. Tissue samples may be taken from animals that have been captured or freshly killed. Non-invasive sources of genetic material include feces, which contains cells sloughed off the lining of the intesti- nal tract (H6éss et al. 1992; Kohn et al. 1995; Wasser et al. 1997). The major problem encoun- tered when amplifying DNA from feces is inhibi- tion of the Tag polymerase (Kohn et al. 1995). Although some problems, such as uneven distribu- tion of cells shed from the intestinal lining (Kohn et al. 1995), have been solved by modifications of techniques (Wasser et al. 1997), others, such as DNA degradation, limit the utility of using feces as a source of DNA (Kohn et al. 1995). A second non- invasive source of DNA is hair (Taberlet and Bouvet 1992; Goosens et al. 1998). Basal cells at the root of a single hair have enough DNA for anal- ysis when used in conjunction with PCR (Higuchi et al. 1988). Compared to feces, hair has the advan- 305 306 tage of being a stable source of DNA, and hair may be removed from live or dead bears (hunter-killed or road-killed). ‘‘Hair-traps” or “hair-snares” have been used by a number of researchers to collect hair samples from bears (Woods et al. 1999). These devices involve a fragrant bait circled by barbed wire with the intent of snagging hair on a barb when a bear goes for the bait. Again, because hair is dry, the DNA sample is stable and hair traps can be checked infrequently. Another source of hair samples that has not been reported to date is wild Crab Apple trees (Malus pumila). Bears climb feral Crab Apple trees in the late summer and early fall to obtain fruit (Bennett et al. 1943; Rogers et al. 1988; Willey 1980), and they leave hairs stuck under thé rough bark or on twigs and broken branches. Trees that have been recently climbed by bears are easy to recognize because the bears’ claws expose orange cambium, and outer branches are frequently broken. The wilted leaves on the broken branches contrast with other leaves on the tree. However, by late autumn the orange color of the cambium layer fades, and without leaves, trees that have been climbed are not easy to spot from a dis- tance. Second growth forests in the northeastern states and provinces have many abandoned apple orchards left from small farms that used to dot the rural land- scape. Apple trees, Lilac bushes (Syringa vulgaris) and old cellar holes are often the only reminders of these former farms. Bear hairs on feral apple trees were not obvious and often required 15—20 min before we were able to find any hairs at all. However, the rate of return per unit effort from tak- ing hairs from apple trees must far exceed the rate of return from hair-traps. During fall 1999 we obtained hair samples from 176 apple trees at 29 locations scattered across Vermont. These samples were col- lected on 21 field days. Hair samples were easiest to find during the period when bears were actively climbing trees, which lasted about four weeks. We were still able to find hairs on trees after this time, but over the next 4—6 weeks bear hairs became pro- gressively more difficult to find as they were blown or washed off of apple trees by wind and rain. Black Bears climb two other trees in the Northeast for fall fruit, Black Cherry (Prunus serotina) and American Beech (Fagus grandifolia) (Bennett et al. 1943; Beeman and Pelton 1980; Rogers et al. 1988; Wolfson 1992). Mature cherry and beech trees are tall and have few side branches on which to catch hairs. Black Cherries have hard rough bark on which it is difficult to see claw marks. Beech trees have smooth bark on which it is easy to see claw marks, but the bark does not have anything to catch hairs. For this reason, apple trees seem unique in being easy for researchers to climb and in having dense prickly branches and rough bark that seem “designed” to capture bear hairs. THE CANADIAN FIELD-NATURALIST. Vol. 116 In summary, apple trees in abandoned orchards are an excellent source of Black Bear hair that can be used for genetic analysis. Discovery of bear hairs requires careful inspection of the trees and frequent climbing to follow the path of bear tracks up the tree. If “wild” apple trees are available, obtaining hair samples in this manner is certainly more efficient than the use of hair-traps, but project objectives would dictate which method is best. Acknowledgments M. Freeman, S. Sweterlitch, C. Scharf, and N. Merrill helped climb apple trees. We thank R. Wells and R. Reay of Vermont Department of Forests and Parks, C. Alexander, D. Blodgett, J. Buck, and T. Decker of Vermont Department of Fish and Wildlife, J. Nelson, S. Williamson, and A. Calfey for informa- tion on locations of Apple trees. Finally, we are especially grateful to L. Orvis for alerting us to the fact that bears were climbing apple trees, as well as for providing information on apple-orchard locations. Literature Cited Alt, G. L., G. L. Matula Jr., F. W. Alt, and J. S. Lindzey. 1980. Dynamics of home range and movements of adult black bears in northeastern Pennsylvania. International Conference on Bear Research and Management 4: 131-136. Beeman, L. E., and M. R. Pelton. 1980. Seasonal foods and feeding ecology of black bears in the Smoky Moun- tains. International Conference on Bear Research and Management 4: 141-147. Bennett, L. J., P. F. English, and R. L. Watts. 1943. The food habits of the black bear in Pennsylvania. Journal of Mammalogy 24: 25-31. Elowe, K. D. 1984. Home range, movements, and habitat preference of black bears (Ursus americanus) in western Massachusetts. M.S. thesis, University of Massachusetts, Amherst. Goosens, B., L. P. Waits, and P. Taberlet. 1998. Plucked hair samples as a source of DNA: reliability of dinu- cleotide microsatellite genotyping. Molecular Ecology 7: 1237-1241. Haig, S. M. 1998. Molecular contributions to conserva- tion. Ecology 79: 413-425. Higuchi, R., C. H. von Beroldingen, G. F. Sensabaugh, and H. A. Erlich. 1988. DNA typing from single hairs. Nature 332: 543-546. Hoss, M., M. Kohn, S. Paabo, F. Knauer, and W. Shroder. 1992. Excrement analysis by PCR. Nature 359) 199. Kohn, M., F. Knauer, A. Stofella, W. Schréder, and S. Padibo. 1995. Conservation genetics of the European brown bear — study using excremental PCR of nuclear and mitochondrial sequences. Molecular Ecology 4: 95-103. Parker, P. G., A. A. Snow, M. D. Shug, C. G. Booton, and P. A. Fuerst. 1998. What molecules can tell us about populations: choosing and using a molecular marker. Ecology 79: 361-382. Paetkau, D., and C. Strobeck. 1994. Microsatellite anal- 2002 ysis of genetic variation in black bear populations. Molecular Ecology 3: 489-495. Pelton, M. R. 2000. Black bear. Pages 389-408 in Ecol- ogy and management of large mammals in North Amer- ica. Edited by S. Demarais and P. R. Krausman. Prentice-Hall, Inc., Upper Saddle River, New Jersey. Rogers, L. L. 1987. Effects of food supply and kinship on social behavior, movements, and population growth of black bears in northeastern Minnesota. Wildlife Monographs 97: 1-72. Rogers, L. L., G. A. Wilker, and A. W. Allen. 1988. Managing northern forests for black bears. Pages 36-42 in Integrating forest management for wildlife and fish. Edited by T. W. Hoekstra and J. Capp. USDA Forest Service, General Technical Report NC-122. Taberlet, P., and J. Bouvet. 1992. Bear conservation genetics. Nature 358: 197. Taberlet, P., H. Mattock, C. Dubois-Paganon, and J. Bouvet. 1993. Sexing free-ranging brown bears Ursus arctos using hair found in the field. Molecular Ecology 2: 399-403. Taberlet, P., and L. P. Waits. 1998. Non-invasive genetic sampling. Trends in Ecology and Evolution 13: 26-27. Taberlet, P., S. Griffin, B. Goosens, et al. 1996. Reliable genotyping of samples with very low DNA NOTES 307 quantities using PCR. Nucleic Acids Research 24: 3189-3194. Waples, R. S. 1989. A generalized approach for estimat- ing effective population size from temporal change in gene frequency. Genetics 121: 379-391. Wasser, S. K., C. S. Houston, G. M. Koehler, G. G. Cadd, and S. R. Fain. 1997. Techniques for application of fecal DNA methods to field studies of Ursids. Molecular Ecology 6: 1091-1097. Whitaker, J. O. Jr., and W. J. Hamilton Jr. 1998. Mammals of the eastern United States, 3rd edition. Cornell University Press, Ithaca, New York. Willey, C. H. 1980. The Vermont black bear. Vermont Fish & Game Department, Montpelier. Wolfson, D. L. 1992. Development of a quantitative pro- cedure to assign a value rating to beech stands as black bear habitat. M.S. thesis, Antioch University, Keene, New Hampshire. Woods, J. G., D. Paetkau, D. Lewis, B. N. McLellan, M. Proctor, and C. Strobeck. 1999. Genetic tagging of free-ranging black and brown bears. Wildlife Society Bulletin 27: 616-627. Received 5 July 2000 Accepted 14 June 2002 Large Clutch Size of a Burrowing Owl, Athene cunicularia, Found in Saskatchewan L. DANIELLE Topp! and JOANN SKILNICK2 ‘Biology Department, University of Regina, Regina, Saskatchewan S4S 0A2 Canada 2Cypress Hills Interprovincial Park, Box 12 Elkwater, Alberta TOJ 1CO Canada Todd, L. Danielle, and Joann Skilnick. 2002. Large clutch size of a Burrowing Owl, Athene cunicularia, found in Saskatchewan. Canadian Field-Naturalist 116(2): 307-308. During the summer of 1999, we observed a pair of Burrowing Owls (Athene cunicularia) lay 14 eggs in a nest near Wilcox, Saskatchewan. This represents the largest clutch size ever recorded for this species. Burrowing Owls typically lay between six and 11 eggs, and prior to our discovery the largest clutch size ever recorded was 12 eggs. Although all 14 eggs hatched, the brood was substantially reduced by cannibalism within two weeks, and in the end only 8 of the 14 chicks fledged. Marked hatching asynchrony, typical in this species, was not observed at this nest. Key Words: Burrowing Owl, Athene cunicularia, clutch size, hatching asynchrony, Saskatchewan. Burrowing Owls (Athene cunicularia) were listed as an endangered species in Canada in 1995 (Wellicome and Haug 1995) due to prolonged and severe declines in the population. This species inhab- its Open prairies, steppes, and deserts throughout North and South America, and reaches the northern limit of its range in Saskatchewan and Alberta. In Canada, Burrowing Owls do not dig their own bur- rows, but rather inhabit burrows originally excavated by animals such as ground squirrels (Spermophilus spp.), Prairie Dogs (Cynomys ludovicianus), and Badgers (Taxidia taxus). Due to their fossorial nesting habits, reliable data on variability in Burrowing Owl clutch size is scarce. Bendire (1892) estimated that clutch sizes ranged from 6 to 11 eggs, with rare instances of 12- egg clutches (Bent 1938). Murray (1976) reported clutch size varying from | to 11 eggs across the species’ North American range. Recently, the use of artificial nest boxes has enabled easier access to nests and thus the ability to collect more accurate data on clutch size. Over a six-year period in Saskatchewan, Wellicome (2000) reported clutch sizes in artificial nest boxes ranging between 6 and 308 12 eggs, with the majority of pairs laying clutches of between 8 and 10 eggs. In the summer of 1999, while monitoring a popula- tion of Burrowing Owls on the Regina Plain, Saskatchewan, we observed a pair of owls lay a clutch of 14 eggs near Wilcox (50° 05’ N, 104° 42’W). To our knowledge, this is the largest clutch ever reported for this species. Boon (1963) reported seeing 14 young Burrowing Owls at a nest site near Elbow, SK, however, that instance was likely a result of juveniles from two or more nests congregating together after fledging (Roy 1996). Intermixing between nests is quite common when juveniles begin to disperse (Todd 2001), and Boon’s observation occurred within the post-fledging dispersal period (31 August). We observed the pair of owls for the first time at their nest burrow on 28 April 1999. We visited the nest once every two to three days as part of an ongo- ing productivity enhancement project (see Wellicome 2000), which involved the installation of artificial nest boxes and the provision of supplemental food to owl pairs after eggs had hatched. We installed an artificial nest box on 2 May 1999; on 5 May 1999 the first egg was laid, and the clutch was complete on 30 May 1999. The first eggs began to hatch on 18 June, and by 21 June all 14 eggs had hatched. Burrowing Owls are known to exhibit hatching asynchrony that often results in more than a week’s difference in the age of the oldest and youngest chicks in the nest (Olenick 1990; Wellicome 2000). Typically, incubation begins after the first few eggs in the clutch are laid, and the female continues to lay eggs at a rate of one every approximately 36 hours until the clutch is complete. This results in an age differential between the oldest (first-hatched) and youngest chicks in the nest, a strategy that often results in the death of the younger siblings from star- vation or cannibalism when food conditions are poor. At this nest, all 14 eggs hatched within three days, resulting in a relatively small age differential within the entire brood. Although all 14 nestlings survived the first few days, within a week three chicks had died, and only 8 of the 14 chicks survived to fledging age (41 days post-hatch). We were unable to determine the ulti- mate cause of death for any of the chicks, or whether it was indeed the youngest chicks that were killed. Chick remains were found inside the box (indicating cannibalism had occurred); however, it is unclear whether the nestlings were killed by their siblings THE CANADIAN FIELD-NATURALIST Vol. 116 and then eaten, or were eaten after dying of some other cause (i.e., starvation, trampling, etc.). Acknowledgments Funding for Burrowing Owl research in Saskatchewan was provided by Wildlife Preservation Trust Canada, World Wildlife Fund and Environment Canada’s Endangered Species Recovery Fund, Canadian Wildlife Service and the Natural Sciences and Engineering Research Council of Canada (NSERC). Logistical support provided by the University of Regina, Saskatchewan Environment and Resource Management (SERM) and Saskatchewan Wetland Conservation Corporation (SWCC). We thank R.G. Poulin, R. M. Brigham and A. J. Erskine for constructive comments on earlier drafts of this manuscript. Documents Cited Wellicome, T.I., and E.A. Haug. 1995. Second update of status report on the Burrowing Owl (Speotyto cunicularia) in Canada. Committee on the Status of Endangered Wildlife in Canada. Canadian Wildlife Service, Ottawa, Ontario. Literature Cited Bendire, C. E. 1892. Life histories of North American birds. U.S. Natural History Museum Special Bulletin Number 1. Bent, A. C. 1938. Life histories of North American birds of prey, part 2. United States National Museum Bulletin 170. Boon, E. 1963. Notes on a Burrowing Owl nest. Blue Jay 21: 104. Murray, G. A. 1976. Geographic variation in the clutch sizes of seven owl species. Auk 93: 602-613. Olenick, B. E. 1990. Breeding biology of Burrowing Owls using artificial nest burrows in southeastern Idaho. M.Sc thesis. Idaho State University, Pocatello, Idaho. 54 pages. Roy, J. F. 1996. Birds of the Elbow. Saskatchewan Natural History Society Special Publication Number 21. Todd, L. D. 2001. Survival and dispersal of juvenile Burrowing Owls (Athene cunicularia) during the post- fledging, pre-migratory period. M.Sc. thesis. University of Regina, Regina, Saskatchewan. 76 pages. Wellicome, T. I. 2000. Effects of food on reproduction in Burrowing Owls (Athene cunicularia) during three stages of the breeding season. Ph.D thesis. University of Alberta, Edmonton, Alberta. 113 pages. Received 11 September 2000 Accepted 3 April 2002 2002 NOTES 309 Dispersal by a Male American Marten, Martes americana DorotTHy M. FECSKE and JONATHAN A. JENKS Department of Wildlife and Fisheries Sciences, Box 2140B, South Dakota State University, Brookings, South Dakota 57007 USA Fecske, Dorothy M., and Jonathan A. Jenks. 2002. Dispersal by a male American Marten, Martes americana. Canadian Field-Naturalist 116(2): 309-311. A radio-collared adult male American Marten in an unharvested population in the Black Hills of South Dakota was initially captured 1 April 1999 and relocated 32 times until found dead 7 December 1999. Over a 21 to 56-day period (5—26 May to 18 April—12 June) it travelled 74 km (straight-line distance between capture site to location at death). The two farthermost locations were 82 km apart. Key Words: American Marten, Martes americana, Black Hills, dispersal, movements Dispersal is the movement of an animal from its natal range to its first or subsequent breeding range, or where it would have reproduced had it survived (Shields 1987). American Martens (Martes ameri- cana) are born in spring (Mead 1994), attain adult size by autumn (Brassard and Bernard 1939), and are able to disperse at that time (Strickland 1994). Dispersal may occur, however, as late as February through the following September (Slough 1989), or possibly, not at all, under conditions of high prey densities (Powell 1994). Transient or resident Martens have dispersed < 20 km from capture sites or previous home ranges (Bateman 1986; De Vos and Gunther 1952). Thompson and Colgan (1987), however, reported longer dispersal distances of three resident adult Martens during years of scarce prey. Two males, ages 3 and 4 years, were trapped 48 and 40 km, respectively, from home ranges they occu- pied for 2 years; and one, five-year-old, female was captured 80 km from its home range of 28 months. We describe long-distance dispersal of an adult male American Marten in an unharvested population in the Black Hills of South Dakota and Wyoming. The Black Hills is an isolated mountain range located in western South Dakota and northeastern Wyoming. The mountains cover about 8400 km?, and are surrounded by the Northern Great Plains (Fecske et al. 2002). The Black Hills has a semi-arid continental and mountain climate. Forests occur at elevations between 1200 and 2100 m. Ponderosa Pine (Pinus ponderosa) forest alliances comprise 84% of the Black Hills forests (Rumble and Anderson 1996); other forest alliances include White Spruce (Picea glauca) and Burr Oak (Quercus macrocarpa) (Hoffman and Alexander 1987). Aver- age annual precipitation ranges from 45 to 66 cm (Orr 1959). Annual mean temperatures range from 5-9° C, with extremes of -40-—44° C (Thilenius 1972). Forests are managed by the United States Department of Agriculture Black Hills National Forest primarily for timber production, livestock grazing, and recreation. Historically, American Martens occurred in South Dakota (Graham and Graham 1994; Turner 1974) and were documented in the Black Hills in the late 1800s and early 1900s (Fredrickson 1993*; Turner 1974). Nonetheless, between 1930 and 1979, no Martens were harvested from the Black Hills. In 1980 and 1981, a reintroduction effort was initiated by the South Dakota Department of Game, Fish and Parks (Fredrickson 1993). Forty-two Martens were released into the northern Black Hills. Additionally, between 1990 and 1993, 83 Martens were released in the central Black Hills. In 1998, a study was initiated to estimate population size and document the current distribution of American Martens within the Black Hills of South Dakota. A male American Marten (Marten number 325) was live-trapped (Tomahawk Live Trap Co., Toma- hawk, Wisconsin) on 1 April 1999 in Black Elk Wilderness Area (Central Black Hills, 103°33'06"N, 43°51'00”W). The Marten was immobilized with ketamine hydrochloride (10.0 mg / 454 g body weight), weighed, aged, and fitted with a radio-collar (Telonics, Inc., Mesa, Arizona). At the time of cap- ture, the animal weighed 1,020 g and was catego- rized as an adult (Poole et al. 1994). We conducted weekly flights from a fixed-wing aircraft to locate the animal. Mean telemetry error was 152.0m (n = 4), which was determined by placing radio-col- lars throughout the study area and having the pilot locate the transmitters. The Marten was not relocated every week due to the limited range of the radio- transmitter, relatively large distances moved between some locations, and inclement weather. We classi- fied the status of the Marten according to that described in Hawley and Newby (1957); Martens that occupied a home range for > 3 months were considered resident, those that occupied a home range > 1 week but < 3 months were considered tem- porary residents, and those present in an area for < | *See Documents Cited section 310 week were transients. We considered a movement greater than 3.6 km indicative of a dispersal event because it surpassed the length of the radius of the mean home-range size (10.2 km?, Zielinski and Kucera 1995) for the species (Kernohan et al. 1994). After the Marten died, age was confirmed using the temporal muscle coalescence (Poole et al. 1994). The estimated age of the animal at the time of its death was > 3 years. Puncture holes in the skull indi- cated that the American Marten was likely killed by a Coyote (Canis latrans) (S. W. Buskirk, University of Wyoming, Laramie, personal communication), and partially consumed. American Marten number 325 was relocated 32 times between the date of capture on | April and the date it was located dead, on 7 December 1999. Initially, on 6, 7, and 17 April 1999, it was located approximately 9.8 km southeast of its capture loca- tion in Black Elk Wilderness Area. Then, on 7 May 1999, the animal was located about 16.2 km west of its capture location. Between 7 and 25 May, the Marten continued traveling north and west, not remaining at any one location for more than one week until it reached the northern Black Hills, between 25 May and 13 June 1999. Over a 21- to 56- day period (5 May—26 May to 18 April—12 June), it traveled 74 km (straight-line distance from the cap- ture site to the location of death); the two farther- most locations were 82 km apart. The Marten was relocated 23 times after it was considered resident (13 September) in the northern Black Hills. The American Marten remained in the northern Black Hills until its death, sometime between 17 November and 7 December 1999. The information obtained on Marten number 325 revealed factors that may influence the American Marten population in the Black Hills. The long dis- tance traveled by the adult Marten could be a result of a fragmented distribution of suitable habitat (Thompson 1994; Soutiere 1979) that occurs in the Black Hills, and the fact that the population may be at carrying capacity (Fecske et al. 2002). The Black Elk Wilderness was one of the reintroduction sites and contains relatively high densities of Martens (Fecske et al. 2002). Adult Martens are known to disperse in high-density, unharvested populations (Davis 1983; Phillips et al. 1998) because of intra- specific competition for limited resources (Powell 1994; Thompson, and Colgan 1987). Acknowledgments This study was supported by Federal Aid to Wildlife Restoration Funds (Study number 7594), administered through the South Dakota Department of Game, Fish and Parks. S. L. Griffin and S. W. Buskirk helped with aging. Laird Flying Service, Ekalaka, Wyoming, and South Dakota Civil Air Patrol provid- ed aerial support. We thank R. T. Bowyer, S. W. THE CANADIAN FIELD-NATURALIST Vol. 116 Buskirk, C. S. DePerno, V.J. Smith, and J. R. Gerard for comments on drafts of this manuscript. Documents Cited (marked * in text) Fredrickson, L. 1993. American Marten reintroduction: a Black Hills success story. Unpublished Report. South Dakota Department of Game, Fish, and Parks, Pierre, South Dakota. 16 pages. Literature Cited Bateman, M. C. 1986. Winter habitat use, food habits and home range size of the Marten, Martes americana, in western Newfoundland. Canadian Field-Naturalist 100: 58-62. Brassard, J. S., and R. Bernard. 1939. Observations on breeding and development of Marten, Martes americana (Kerr). Canadian Field-Naturalist 53: 15-21. Davis, M. H. 1983. Post-release movements of introduced Marten. Journal of Wildlife Management 47: 59-66. De Vos, A., and S. E. Gunther. 1952. Preliminary live- trapping studies of Marten. Journal of Wildlife Manage- ment 16: 207-214. Fecske, D. M., J. A. Jenks, and V. J. Smith. 2002. Field evaluation of a habitat-relation model for the American marten. The Wildlife Society Bulletin 30: 775-782. Graham, R. W., and M. A. Graham. 1994. Late Qua- ternary distribution of Martes in North America. Pages 26-58 in Martens, sables, and fishers: biology and con- servation. Edited by S. W. Buskirk, A. S. Harestead, M. G. Raphael, and R. A. Powell. Comstock Publishing Associates, Ithaca, New York. 484 pages. Hawley, V. D., and F. E. Newby. 1957. Marten home ranges and population fluctuations. Journal of Mam- malogy 38: 174-184. Hoffman, G. R., and R. R. Alexander. 1987. Forest vege- tation in the Black Hills National Forest of South Dakota and Wyoming: a habitat type classification. USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, Research Paper RM-276, Fort Collins, Colorado, USA. Kernohan, B. J., J. A. Jenks, and D. E. Naugle. 1994. Movement patterns of white-tailed deer at Sand Lake National Wildlife Refuge, South Dakota. Prairie Natur- alist 26: 293-300. Mead, R. A. 1994. Reproduction in Martes. Pages 404— 422 in Martens, sables, and fishers: biology and conser- vation. Edited by S. W. Buskirk, A. S. Harestead, M. G. Raphael, and R. A. Powell. Comstock Publishing Asso- ciates, Ithaca, New York. 484 pages. Orr, H. K. 1959. Precipitation and streamflow in the Black Hills. USDA Station Paper 44. Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado. 25 pages. Poole, K. G., G. M. Matson, M. A. Strickland, A. J. Magoun, R. P. Graf, and L. M. Dix. 1994. Age and sex determination for American Martens and fishers. Pages 204-223 in Martens, sables, and fishers: biology and conservation. Edited by S. W. Buskirk, A.S. Harestead, M. G. Raphael, and R. A. Powell. Comstock Publishing Associates, Ithaca, New York. 484 pages. Phillips, D. M., D. J. Harrison, and D. C. Payer. 1998. Seasonal changes in home-range area and fidelity of Martens. Journal of Mammalogy 79: 180-190. 2002 Powell, R. A. 1994. Structure and spacing of Martes pop- ulations. Pages 101-121 in Martens, sables, and fishers: biology and conservation. Edited by S. W. Buskirk, A. S. Harestead, M. G. Raphael, and R. A. Powell. Comstock Publishing Associates, Ithaca, New York. 484 pages. Rumble, M. A., and S. H. Anderson. 1996. Micro- habitats of Merriam’s turkeys in the Black Hills, South Dakota. Ecological Applications 6: 326-334. Shields, W. M. 1987. Dispersal and mating systems: inves- tigating their causal connections. Pages 3-24 in Mammalian dispersal patterns: the effects of social struc- ture on population genetics. Edited by B. D. Chepko-Sade and Z. T. Halpin. The University of Chicago Press, Chicago, Illinois. 342 pages. Slough, B. G. 1989. Movements and habitat use by trans- planted Marten in the Yukon territory. Journal of Wildlife Management 53:991—997. Soutiere, E.C. 1979. Effects of timber harvesting on Marten in Maine. Journal of Wildlife Management 43: 850-860. Strickland, M. A. 1994. Harvest management of fishers and American Martens. Pages 149-164 in Martens, sables, and fishers: biology and conservation. Edited by S. W. Buskirk, A. S. Harestead, M. G. Raphael, and R. NOTES 311 A. Powell. Comstock Publishing Associates, Ithaca, New York. 484 pages. Thilenius, J. F. 1972. Classification of deer habitat in the ponderosa pine forest of the Black Hills, South Dakota. U.S. Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado, USA. Thompson, I. D. 1994. Marten populations in uncut and logged boreal forests in Ontario. Journal of Wildlife Management 58: 272-280. Thompson, I. D., and P. W. Colgan. 1987. Numerical responses of Martens to a food shortage in northcentral Ontario. Journal of Wildlife Management 51: 824-835. Turner, R.W. 1974. Mammals of the Black Hills of South Dakota and Wyoming. Miscellaneous Publication No. 60, University of Kansas Publications, Museum of Natural History. 178 pages. Zielinski, W. J., and T.E. Kucera. 1995. American Marten, fisher, lynx, and wolverine: survey methods for their detection. General Technical Report psw-gtr 157, US Forest Service, Albany, California. 163 pages. Received 27 December 2000 Accepted 7 May 2002 Territorial Marking by Lone Male Gray Wolves, Canis lupus RONALD N. SCHULTZ! and PAMELA C. WILSON2 'Wisconsin Department of Natural Resources, 8770 Hwy J, Woodruff, Wisconsin 54568 USA 2University of Kansas, Lawrence, Kansas 66045-2106 USA Schultz, Ronald N., and Pamela C. Wilson. 2002. Territorial marking by lone male Gray Wolves, Canis lupus. Canadian Field-Naturalist 116(2): 311-313. We documented three separate instances in which alpha male Gray Wolves (Canis lupus) scent marked and remained in their original pack’s territory after all the other members of the original pack were gone due to death or dispersal. They continued to scent mark until they attracted another mate to the territory and began a new pack. Key Words: Gray Wolf, Canis lupus, pack, alpha, sent marking, lone, territory, Two Toes, RLUs. Most previous literature on the behavior of solitary Gray Wolves has focused on single yearling or adult females remaining in their original pack territory to attract a new mate (Ballard et al. 1987), and dispersing offspring of alpha pairs (Mech 1973; Van Ballenberghe 1983; Messier 1985; Gese and Mech 1991) with occa- sional mention of “peripheral” wolves that inhabit the edges of pack territories (Mech 1970). It was often assumed that solitary male wolves do not maintain ter- ritories by scent marking, but disperse and attempt to colonize vacant habitat or join existing packs (Ballard et al. 1987). Implicit in this assumption is that alpha males left alone will not maintain and scent mark terri- tories previously occupied and maintained by their entire pack (Rothman and Mech 1979: 753). Three adult male wolves from three different packs were monitored from 1991-1995, in north central Wisconsin. Two of the males, M128 and M201, and several of their pack members were radio-collared and located by aerial or ground surveillance once or twice a week. Aerial telemetry was used to delineate territories (determined by > 35 winter and > 25 summer locations (Fuller and Snow 1988)) using minimum area polygon criteria (Mohr 1947). Visual observations, winter snow track sur- veys (Thiel 1978; Thiel and Welch 1981) and sum- mer howling surveys (Harrington and Mech 1982) were used to determine whether wolves were alone or with other pack members. During winter tracking, raised leg urinations (RLUs) were used to determine the presence of an alpha wolf, and double RLUs (with estrus blood) were used to determine the pres- ence of a breeding pair (Peters and Mech 1975; Rothman and Mech 1979). ¥ 312 THE CANADIAN FIELD-NATURALIST M128, an adult male of the Bootjack Lake Pack, was captured and collared on 23 July 1991. Sixty-six aerial-radio locations of an adult female (F099) had been used to delineate the territory inhabited by this pack during 1987-1988. There was evidence that M128 inhabited the area with other pack members immediately prior to his capture. Two wolves responded to howling on 13 and 18 July 1991, and previous winter tracking surveys (1990-91) indicat- ed there had been three wolves. Data obtained from 52 aerial locations, four howling surveys, three sightings and two snow tracking surveys indicated that wolf M128 remained alone in his territory from the time of capture until 3 February 1992. Two RLUs found on a winter track survey on 6 January 1992 indicated he continued to scent mark his terri- tory without a mate or other pack members. Wolf M128 was first seen with another wolf on 3 February 1992. The next day tracks of two animals (one being M128) and double RLUs with estrus blood were found, indicating that M128 had been joined by an adult female. He was seen on 17 subsequent aerial locations with one or more wolves during 6 February 1992-12 April 1993. Our second case of a jone wolf scent marking and maintaining a territory involved wolf M201, an adult male from the Little Rice River Pack. M201 was captured and radio collared 23 October 1991. From capture until 31 August 1993, he occupied a territory with one or more pack members, based on aerial observations, track surveys, and the presence of RLUs with estrus blood. From 1 September 1993 until 12 March 1995, M201 was alone and remained within his territory based on 71 telemetry locations. During this period, wolf M201 continued to scent mark. M201 was snow tracked a total of 5.8 km, on 24 February 1994, and 31 January 1995. We found no tracks of other wolves but RLUs were found in three different locations on these track surveys. Six aerial observations also indicated he was alone. Three RLUs, one with estrus blood, were observed on 12 March 1995 while we were following tracks of two wolves (one being M201) for 1.9 km in M201’s territory. This was the first evidence of a second wolf since | September 1993. Our third case involved an un-collared male wolf nicknamed Two Toes. He was missing the center two toes on one front foot, making his tracks easy to identify. Two Toes appeared to be the alpha male of the Ranger Island Pack in January 1992. Two wolves, one of which was missing two toes, was tracked a total of 4.8 km in that territory on 14 January and 31 March 1992. Evidence of three RLUs, with two containing estrus blood, indicated a breeding pair in this pack. No wolf tracks other than those of Two Toes were found on track surveys con- ducted on 8 December 1993, 22 January 1994, and 17 February 1994. We found six RLUs on snow Vol. 116 banks along four different gravel roads in the Ranger Islands Pack’s territory. Two of the RLUs were deposited on snow clumps located at road intersec- tions normally scent marked by this pack. This indi- cated he continued to mark his territory while alone. Tracks of two wolves one of which was missing two toes, were followed 6.7 km on | January 1995 in the southwest corner of the Ranger Island Pack’s territo- ry. It appeared that “Two Toes” may: have found a new mate after > 2 years of maintaining this territory alone. The three males mentioned here scent marked an average of 0.4 RLUs per km on main roads while alone in their territories. This was less than when they were with mates or other pack members (0.7 RLUs per km). It seems that maintaining a territory after the death or dispersal of all other pack members is a viable option for some lone male wolves (Rothman and Mech 1979) and apparently lone female wolves (Ballerd et al. 1987). It could be assumed that when a suitable territory is established, the best strategy may be to scent mark this territory and wait for a mate, instead of venturing out in unfa- muiliar territory to find one. Acknowledgments Persons assisting in monitoring wolves M201, M128, M107, M101, F099, and M104 included Ray Marvin, Daniel Doberstein and Gary Peterson. Wolf monitoring was funded by U.S. Fish and Wildlife Service, Section 6 Grant, Wisconsin Endangered Species Fund. We are grateful to David Seibel for his assistance in using graphic software and for pro- viding his editorial skills. Special thanks to Sheri Buller, Bruce Kohn, David Mech, Adrian Wydeven and Linda Winn for their help in editing. Literature Cited Ballard, W.B., J.S. Whitman, and C. L. Gardner. 1987. Ecology of an exploited wolf population in south- central Alaska. Wildlife Monograph 98. 54 pages. Fuller, T. K., and W. J. Snow. 1988. Estimated wolf den- sities from radio telemetry data. Wildlife Society Bulletin 16: 367-370. Gese, E. M., and L. D. Mech. 1991. Dispersal of wolves (Canis lupus) in northeastern Minnesota, 1969-1989. Canadian Journal of Zoology 69: 2946-2955. Herrington, F. H., and L. D. Mech. 1982. An analysis of howling response parameters useful for wolf pack cen- susing. Journal of Wildlife Management 46: 686-693. Mech, L. D. 1970. The wolf: the ecology and behavior of an endangered species. Doubleday, New York, N.Y. 389 pages. Mech, L. D. 1973. Wolf numbers in the Superior National Forest of Minnesota. USDA Forest Service Research Paper NC-97. North Central Forest Experimental Station, U.S. Forest Service. St. Paul, Minnesota. Messier, F. 1985. Solitary living and extraterritorial movements of wolves in relation to social status and prey abundance. Canadian Journal of Zoology 63: 239-245. 2002 NOTES 313 Mohr, C.O. 1947. Table of equivalent populations of North American small mammals. American Midland Naturalist 37: 223-249. Peters, R. P., and L. D. Mech. 1975. Scent-marking in wolves. American Scientist 63: 628-637. Rothman, R. J., and L. D. Mech. 1979. Scent-marking in lone wolves and newly formed pairs. Animal Behaviour 27: 750-760. Received 19 July 2000 Thiel, R. P., and R. J. Welch. 1981. Evidence of recent Accepted 21 April 2002 breeding activity in Wisconsin wolves. American Midland Naturalist 106: 401-402. Van Ballenberghe, V. 1983. Extraterritorial movements and dispersal of wolves in south central Alaska. Journal of Mammalogy 64: 168-171. Estimating the Weight of Wolves, Canis lupus, from Chest Girth Measurements JOHN P. HART! and DAvipD H. JAMIESON? 1United States Department of Agriculture, Wildlife Services, 34912 U.S. Highway 2, Grand Rapids, Minnesota 55744 USA 2Arkansas State University-Newport Campus, Department of Biological Sciences, 7648 Victory Boulevard, Newport, Arkansas 72112 USA Hart, John P., and David H. Jamieson. 2002. Estimating the weight of Wolves, Canis lupus, from chest girth measure- ments. Canadian Field-Naturalist 116(2): 313-314. Regression equations relating weight with chest girth have been reported for a variety of wildlife species, but not Wolves (Canis lupus). We recorded the weights and chest girths of 114 Wolves taken in central and northern Minnesota between 1995 and 1997. Regression equations were developed for the estimation of Wolf weight from chest girth. A significant cor- relation existed between body weight and chest girth (7? = 0.815, n = 114, P < 0.001). There was no significant difference between regression equations when grouped by sex. Key Words: Wolf, Canis lupus, chest girth, Minnesota. Because weights are indicators of physical condi- tion (Kirkpatrick 1980) they are useful in research and management of wildlife species and are often obtained from harvested Wolves (Canis lupus), or those captured for research purposes. Several studies have demonstrated that weights can be accurately predicted from chest girth measure- ments of a variety of mammals. Payne (1976) devel- oped an equation that would predict the weight of Black Bears (Ursus americanus) within about 95 per- cent of the true value for Newfoundland. Likewise, relationships between weight and chest girth have been established for Caribou (Rangifer tarandus), (McEwan and Wood 1966), White-tailed Deer (Odocoileus virginianus) (Urbston et al. 1976), Bison (Bison bison) (Kelsall et al. 1978), Mountain Goats (Oreamnos americanus) (Rideout and Worthen 1975), Grizzly Bears (Ursus arctos) (Nagy et al. 1984), Polar Bears (Ursus maritimus) (Kolenosky et al. 1989), Elk (Cervus elaphus) (Millspaugh and Brundige 1996) as well as several East African mammals (Talbot and McCulloch 1965), to name a few. No such relation- ship has been reported for the Wolf. This study was conducted to determine if a correlation between Wolf weight and chest girth existed. Methods Wolves were captured with foothold traps and neck snares during routine livestock depredation control activities by U.S. Department of Agriculture — Wildlife Services personnel. Wolves were taken in 15 counties in northern and central Minnesota from June Chest Girth/Weight Correlation (Males and Females n=114) 50 + | 40 + DB = 30+ E 2 20 + = 10 + | dis — Sk : - | 25 45 65 eo . 0 | y = 0.8916x - 30.59 R*= 0.8154 | Chest Girth (cm) | FIGURE |. Relationship between chest girth (cm) and weight (kg) of Wolves in Minnesota, 1995-1997. em ee ee eC ee Ce” 314 1995 through October 1997. Wolves were weighed to the nearest kilogram using a spring scale. Chest girths were taken in centimeters with a standard flexible measuring tape while the Wolves were lying on their sides; chest girth measurements were taken at the largest body circumference behind the front leg but anterior to the xyphoid process. Care was taken to make sure the tape was snug but not tight. All weights and measurements were taken post-mortem. We used Microsoft Excel 97 and Minitab 2000 version 13 to determine regression equations. Results and Discussion Weight and chest girth measurements were taken from 114 wolves (53 males and 61 females). A sig- nificant correlation existed between weight and chest girth for all Wolves (7? = 0.815, P < 0.001, n = 114, y = 0.8916x-30.59, where y = estimated whole body weight in kg and x = chest girth in cm; Figure 1). Correlations existed for both male and ‘female Wolves as separate groups (r? = 0.816, n = 53, P<0.001 for males, 72 = 0.737, n= 61, P <0:001 for females). We conducted a “test for parallelism” and a “test for intercepts” to determine if the slopes of the two regression lines were significantly differ- ent. No significant difference was found. Increments in girth size, whether male or female, tended to pro- duce like increases in weight. In addition, the regres- sion lines were not just parallel, they were virtually identical. Acknowledgments Support for this project was provided by Pam Cicerello, Pulaski Technical College, Bob Stiger, Arkansas State University — Newport and Eugene McKay, Arkansas State University — Beebe. W. J. Paul, U.S. Department of Agriculture — Wildlife Services assisted in procurements of wolf measure- ments, Thomas Bishop and Roger Abernathy, Arkansas State University — Jonesboro assisted THE CANADIAN FIELD-NATURALIST Vol. 116 with statistical analysis, and L. D. Mech, U.S. Geo- logical Survey, reviewed this manuscript. Literature Cited Kelsall, J. P., E. S. Telfer, and M. Kingsley. 1978. Relationship of bison weight to chest girth. Journal of Wildlife Management 42: 659-661. Kirkpatrick, R. L. 1980. Physiological indices in wildlife management. Pages 99-112 in Wildlife management techniques manual. 4th edition. Edited by S. D. Schemnitz. The Wildlife Society, Washington D.C. 686 pages. Kolenosky, G. B., N. J. Lunn, C. J. Greenwood, and K. F. Abraham. 1989. Estimating the weight of polar bears from body measurements. Journal of Wildlife Management 53: 188-190. McEwan, E. H., and A. J. Wood. 1966. Growth and development of the barren ground caribou. I. Heart girth, hind foot length, and body weight relationships. Can- adian Journal of Zoology 44: 401-411. Millspaugh, J. J., and G. C. Brundige. 1976. Estimating elk weight from chest girth. Wildlife Society Bulletin 24: 58-61. Nagy, J. A., M. C. Kingsley, R. H. Russell, A. M. Pearson, and B. C. Goski. 1984. Relationship of weight to chest girth in the grizzly bear. Journal of Wildlife Management 48: 1439-1440. Payne, N. F. 1976. Estimating live weight of black bears from chest girth measurements. Journal of Wildlife Management 40: 167-169. Rideout, C. B., and G. L. Worthen. 1975. Use of girth measurement for estimating weight of mountain goats. Journal of Wildlife Management 39: 705-708. Talbot, L. M., and J. S.G. McCulloch. 1965. Weight estimations for East African mammals from body mea- surements. Journal of Wildlife Management 29: 84-89. Urbston, D. F., C. W. Smart, and P. F. Scanlon. 1976. Relationship between body weight and heart girth in white-tailed deer from South Carolina. Proceedings Annual Conference Southeast Association Fish and Wildlife Agencies 30: 471-473. Received 9 October 2000 Accepted 29 June 2002 2002 NOTES 315 Differential Use of a Wolf, Canis lupus, Pack Territory Edge and Core L. DAvip MEcu! and ELIZABETH K. HARPER? ‘Biological Resources Division, U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 — 37" St., SE, Jamestown, North Dakota 58401-7317 USA International Wolf Center, 3300 Bass L. Road, #202, Minneapolis, Minnesota 55429 USA Mailing address: The Raptor Center, University of Minnesota, 1920 Finch Avenue, St. Paul, Minnesota 55108 USA Mech, L. David, and Elizabeth K. Harper. 2002. Differential use of a Wolf, Canis lupus, pack territory edge and core. Canadian Field-Naturalist 116(2): 315-316. Based on 418 radio-locations of a Minnesota Wolf pack, Wolves were found at significantly fewer locations per area in the outer 2 km of the territory than in the core. This finding supports an hypothesis that buffer zones exist between pack terri- tories and may explain why prey survive longer there. Key Words: Wolf, Canis lupus, buffer zone, territory, White-tailed Deer, Odocoileus virginianus, predator, prey Mech (1977a,b) postulated that Wolves (Canis lupus) may use the edges of their territories less than their centers. This proposal was based on his finding that the main prey of the wolves in his northeastern Minnesota study area, White-tailed Deer (Odo- coileus virginianus), survived longer along Wolf pack territory edges than centers (Hoskinson and Mech 1976, Mech 1977b, Rogers et al. 1980, Nelson and Mech 1981). Similar Wolf-deer relationships were observed in northwestern Minnesota (Fritts and Mech 1981) and on Vancouver Island, British Columbia (Hebert et al. 1982, Hatter 1984). Furthermore, theoreticians have found mathematical support for the pack territory edge as a prey refuge (Lewis and Murray 1993), and others have described similar prey-rich zones between warring Indian tribes (Hickerson 1965, 1970, Martin and Szuter 1999). The explanation proposed for higher deer survival along Wolf pack territory edges was the possibility that Wolves spend less time along the edges because of the increased chance of encountering neighboring packs and hence intraspecific strife there. The terri- tory edges were thus thought to be a “no man’s land” or buffer zone between neighboring packs. Indeed a disproportionate number of Wolf-killed Wolves have been found in these buffer zones both in Minnesota (Mech 1994) and Alaska (Mech et al. 1998). Although Mech (1977b) stated that the pack he studied seemed to spend less time in its territory buffer zone, only Carbyn (1983) has analyzed data to test this hypothesis. Such a test requires more loca- tion data for a single pack than are usually obtained. Here we examine a large collection of location data for one pack. We hypothesized that if Wolves tend to spend less time in their territory buffer zone, the number of Wolf locations determined by telemetry there should be significantly less per area than in the remaining core area. Study area and Methods We studied the Harris Lake (HL) Wolf pack that inhabited the Superior National Forest (SNF) of northeastern Minnesota. The Wolf packs in the SNF are spaced into territories, and the HL Pack territory, 19.2 km southeast of Ely (48°N, 92°W.), was sur- rounded by five or six other pack territories (Mech 1973). The terrain, vegetation, and land use in the territory are typical of the surrounding region and were described by Mech (1977b). The HL pack data we used were collected by aerially radio tracking eight members of the pack of two to nine Wolves from late 1968 through early 1975 (Mech 1977b). We drew a minimum convex polygon (MCP; Mohr 1947) around a plot of the composite data from this period (Mech 1977b: Figure 5N). We divided the plot into a core area and a peripheral zone separated by a line parallel- ing the MCP 2 km inside (Mech 1977a). The areas of the peripheral zone and core were calculated, and the number of locations in each were tallied. Multiple points at the same location were only counted once; because most such multiple points were in and near dens in the core, any resulting bias from deleting these points would weigh against the hypothesis of fewer locations in the peripheral zone. Results Some 1,192 Wolf data points were obtained, and of these, 457 involved more than | radioed Wolf together, leaving 735 individual data points (Mech 1977b). Of these 735 points, 418 were at different locations. Of the 418 locations, 102 lay in the peripheral zone, and 316 in the core. The peripheral zone comprised 85 km”, and the core, 138 km?. Thus the density of locations at which radioed wolves were found was 1.2 locations/km? in the periphery, and 2.3 locations/km? in the core, a significant dif- ference (X?= 18.10; P < 0.001; d.f. = 1). Conclusion Except for data from one of two packs studied by Carbyn (1983), these results represent the first sup- eee ——— ee eo + ee 316 port for the hypothesis that Wolves spend less time along the edge of their territory than in the center (Mech 1977a,b), and this finding may explain why deer survive longer along those edges, as document- ed above. This analysis, however, involves only a single Wolf pack territory. Additional testing with other territories for which large amounts of location data are available is necessary to further test this hypothesis further. Acknowledgments This study was supported by the Biological Resources Division of U.S. Geological Survey and the U.S. Department of Agriculture, North Central Research Station. Literature Cited Carbyn, L. N. 1983. Wolf predation on elk in Riding Mountain National Park, Manitoba, Journal a Wildlife Management 47: 963-976. Fritts, S. H., and L. D. Mech. 1981. Dynamics, move- ments, a feeding ecology of a newly protected wolf population in northwestern Minnesota. Wildlife Mono- graphs 80: 1-79. Hatter, I. W. 1984. Effects of wolf predation on recruit- ment of black-tailed deer on northeastern Vancouver Island. Master’s thesis, University of Idaho, Moscow. Hebert, D. M., J. Youds, R. Davies, H. Langin, D. Janz, and G. W. Smith. 1982. Preliminary investigations of the Vancouver Island wolf (Canis lupus crassodon) prey relationships. Pages 54—70 in Wolves of the World. Edited by F. H. Harrington and P. C. Paquet. Noyes Publishers, Park Ridge, New Jersey. Hickerson, H. 1965. The Virginia deer and intertribal buffer zones in the upper Mississippi Valley. Pages 43-66 in Man’s culture and animals. Publication 8. Edited by A. Leeds and A. P. Vayda. American Association Advancement of Science, Washington, D.C. Hickerson, H. 1970. The Chippewa and their neighbors: a THE CANADIAN FIELD-NATURALIST Vol. 116 study in ethnohistory. Holt, Rinehart and Winston, New York. Hoskinson, R. L., and L. D. Mech. 1976. White-tailed deer migration and its role in wolf predation. Journal of Wildlife Management 40: 429-441. Lewis, M. A., and J. D. Murray. 1993. Modeling territo- riality and wolf-deer interactions. Nature 366: 738-740. Martin, P. S., and C. R. Szuter. 1999. War zones and game sinks in Lewis and Clark’s west. Conservation Biology 13: 36-45. Mech, L. D. 1973. Wolf numbers in the Supeniet National Forest of Minnesota. USDA Forest Service Research Paper NC-97. North Central Forest Experiment Station, St. Paul, Minnesota. Mech, L. D. 1977a. Wolf-pack buffer zones as prey reser- voirs. Science 198: 320-321. Mech, L. D. 1977b. Population trend and winter deer con- sumption in a Minnesota wolf pack. Pages 55-83 in Proceedings of the 1975 Predator Symposium. Edited by R. L. Phillips and C. Jonkel. Montana Forest and Conservation Experiment Station. University of Montana, Missoula. Mech, L. D. 1994. Buffer zones of territories of gray wolves as regions of intraspecific strife. Journal of Mammalogy 75: 199-202. Mech, L. D., L. G. Adams, T. J. Meier, J. W. Burch, and B. W. Dale. 1998. The wolves of Denali. University of Minnesota Press, Minneapolis. Mohr, C.O. 1947. Table of equivalent populations of North American small mammals. American Midland Naturalist 37: 223-249. Nelson, M. E., and L. D. Mech. 1981. Deer social organi- zation and wolf predation in northeastern Minnesota. Wildlife Monographs 77: 1-53. Rogers, L. L., L. D. Mech, D. K. Dawson, J. M. Peek, and M. Korb. 1980. Deer distribution in relation to wolf pack territory edges. Journal of Wildlife Management 44: 253-258. Received 22 January 2001 Accepted 3 April 2002 2002 NOTES 317 Extension de I|’aire de distribution de la Salamandre a quatre doigts, Hemidactylium scutatum, dans |’ est du Québec, et notes sur |’ habitat. JEAN-FRANCOIS DESROCHES! et BENOIT COUTURE? 18940 avenue Pradier, Charlesbourg, Québec G1G 5S5 Canada 2380 route 143, Windsor, Québec J1S 2X2 Canada Desroches, Jean-Frangois, et Benoit Couture. 2002. Extension de l’aire de distribution de la Salamandre 4 quatre doigts, Hemidactylium scutatum, dans |’ est du Québec, et notes sur |’habitat. Canadian Field-Naturalist 116(2): 317-318. Une Salamandre 4 quatre doigts juvénile a été trouvée au Lac du Castor, dans la municipalité de Saint-Victor-de-Tring (46°6'19”N, 70°50’55”0 ) le 31 juillet 2000. Cette mention constitue la premiére observation de l’espéce dans la région de Chaudiére-Appalaches et se retrouve approximativement a 160 kilométres au nord-est et 4 145 kilométres au sud-est des deux sites les plus prés d’ou l’espéce est connue. Le milieu présent a cet endroit n’est pas une tourbiére 4 sphaigne, habitat cité le plus souvent au Québec pour cette espéce, mais plutdt un étang a castors en milieu forestier avec secteurs herbeux. Mots-clés : Salamandre 4 quatre doigts, Hemidactylium scutatum, aire de distribution, sphaigne, habitat, Québec. La salamandre 4 quatre doigts (Hemidactylium scutatum) est rare au Québec (Bider et Matte 1994) et figure sur la liste des espéces susceptibles d’étre désignées menacées ou vulnérables (Beaulieu 1992). A cause de ses meeurs discrétes et des habitats qu’elle fréquente, tels les tourbiéres a sphaigne (Sphagnum sp.), cette salamandre est tres difficile a trouver (Bonin 1998; Bider et Matte 1994). Un spécimen a été trouvé au Lac du Castor, municipalité de Saint-Victor-de-Tring (46°6'19"N, 70°50’55"0), Québec, le 31 juillet 2000. Cette mention se retrouve a approximativement 160 kilométres au nord-est (tourbiére de Marlington, comté de Stanstead, @ucbec, 45°2'N, 72°11’ O: Sharbel 1990) et 145 kilométres au sud-est (Saint-Maurice, comté de Champlain, Québec, 46°25'N, 72°30’O: Bider et Matte 1991) des sites les plus prés ou l’espéce a déja été observée, qui se trouvent dans d’autres bassins versants éloignés. Le spécimen trouvé au Lac du Castor est un juvénile (longueur totale : 49,5 mm; longueur mu- seau-cloaque : 26,0 mm) et il a été découvert sous une planche de fibre de bois pressée de 70cm X 50 cm, sous laquelle se trouvaient d’autres planches semblables. Des photographies de la salamandre et de habitat ont été prises par les auteurs et le spéci- men a été déposé dans la amphibien et reptile Musée canadien de la nature (CMAR 35371). Le Lac du Castor n’est pas une tourbiére 4 sphaigne, habitat souvent décrit pour cette espéce (Bonin 1998; Conant et Collins 1998; Behler et King 1996; Bider et Matte 1994; Cook 1984). Il s’agit d’un étang a castors en milieu forestier, entouré de rives d’herbacées et de Myrique baumier (Myrica gale) outre des secteurs a quenouilles (Typha sp.). La Salamandre a quatre doigts était 4 quatre (4) métres du bord de |’étang, dans un secteur herbeux ouvert avec roches au sol, pres de |’ émissaire. Le site a été visité dans un rayon d’environ 200 métres afin de le décrire et de vérifier la présence de sphaigne. Le secteur de |’étang a proximité présente des buttons herbeux et des arbres morts et aucune sphaigne n’y a été vue. Quant a |’émissaire, il inonde en partie le milieu forestier adjacent constitué de forét mixte ol poussent le Sapin baumier (Abies bal- samea), le Bouleau gris (Betula populifolia), le Bouleau jaune (Betula alleghaniensis), le Bouleau blanc (Betula papyrifera), ’Erable rouge (Acer rubrum) et le Thuya occidental (Thuja occidentalis). Les rives de cette zone inondée sont parfois her- beuses et quelques ilots de graminées sont aussi présents a quelques endroits. Le sol forestier en bor- dure de |’émissaire présente une litiére de feuilles mortes assez séches et d’aiguilles de coniféres. La microtopographie est légérement accidentée et on note la présence de quelques biches avec mousse. Aucune sphaigne n’a été vue. Bien que les milieux ot 1’on retrouve de la sphaigne soient les habitats les plus souvent cités pour l’espéce au Québec (Bonin 1998; Bider et Matte 1991; Sharbel 1990), certains auteurs y ont découvert cette salamandre en des endroits ot la sphaigne était limitée a de petites parcelles disper- sées (Gordon 1979; Denman 1965). La sphaigne constituerait le microhabitat optimal pour la ponte des ceufs mais ne serait pas obligatoire (Petranka 1998). D’ ailleurs, plusieurs observations de l’espéce faites dans des localités périphériques aux Etats-Unis sont typiquement dans les pentes boisées ou trés peu mentionnent la présence de sphaigne (Dundee 1968). La Salamandre a quatre doigts effectue des migra- tions et les juvéniles quittent les sites de reproduc- tion quelques semaines aprés leur métamorphose pour aller dans les foréts environnantes, ou ils demeureront jusqu’a leur maturité sexuelle (Petranka 1998). Il est possible que la sphaigne soit présente en dehors du rayon de 200 m inventorié au Lac du Castor et que la salamandre, un juvénile, s’en soit éloigné. Il se pourrait également qu’a cet endroit la Salamandre a quatre doigts ponde dans d’autres 318 types de substrats. Outre la sphaigne, on rapporte des pontes de l’espeéce dans divers types de mousses, dans la litiére de feuilles mortes, sous de l’écorce au sol, dans le bois en décomposition et dans des touffes d’herbe (Petranka 1998). Bishop (1941) men- tionne des observations faites dans des bois ouverts ou l’herbe pousse et sous des planches et biches prés d’un étang. Ces descriptions concordent avec |’ habi- tat retrouvé au Lac du Castor. D’ autres recherches permettront peut-étre de déterminer si la présence de la salamandre a quatre doigts dans ce type d’habitat constitue un fait rare au Québec ou si l’espéce s’y retrouve plus souvent que ce que |’on croit. II serait également intéressant de chercher ot pondent les Salamandres a quatre doigts dans ce secteur. Remerciements Les auteurs remercient M. Guy Leclerc, proprié- taire du site, pour en avoir permis l’accés, de méme que M. Joél Bonin qui a bien voulu réviser et criti- quer l’article avant soumission. Ils tiennent égale- ment a remercier M. Stephen J. Darbyshire, de méme qu’un correcteur anonyme pour leurs com- mentaires et suggestions fort pertinents lors de la révision du texte. Merci finalement 4 Mme Michéle Steigerwald, responsable adjointe de la collection d’amphibiens et reptiles du Musé canadien de la nature, pour sa précieuse collaboration lors du pro- cessus d’intégration du specimen dus la collection du MCN. Littérature citée Beaulieu, H. 1992. Liste des espéces de la faune vertébrée susceptibles d’étre désignées menacées ou vulnérables. Ministére du Loisir, de la Chasse et de la Péche. Québec. 107 pages. Behler, J. L., et F. W. King. 1996. Field guide to North American reptiles and amphibians. National Audubon Society. Alfred A. Knopf Inc. U.S.A. 743 pages. THE CANADIAN FIELD-NATURALIST Vol. 116 Bider, J.-R., et S. Matte. 1994. Atlas des amphibiens et reptiles du Québec. Société d’histoire naturelle de la val- lée du Saint-Laurent et ministére de l’Environnement et de la Faune du Québec, Direction de la faune et des habitats. Québec. 106 pages. Bider, J.-R., et S. Matte. 1991. Atlas des amphibiens et reptiles du Québec 1988—1989-1990, version détaillée. Société d’ histoire naturelle de la vallée du Saint-Laurent et ministére du Loisir, de la Chasse et de la Péche du Québec. Québec. 429 p. Bishop, S.C. 1941. The salamanders of New York. The New York State Museum Bulletin 324 : 1-365. Bonin, J. 1998. Status report on the Four-toed Salaman- der Hemidactylium scutatum in Canada. Rapport préparé pour le COSEWIC (COSEPAC, Comité sur la situation des espéces en péril au Canada). 28 pages. Conant, R., et J. T. Collins. 1998. A field guide to rep- tiles and amphibians of eastern and central North America, 3rd edition, expanded. Houghton Mifflin Com- pany, U.S.A. 616 pages. Cook, F. R. 1984. Introduction aux amphibiens et reptiles du Canada. Musée national des sciences naturelles, Musées nationaux du Canada. Ottawa, Canada. 211 p. Denman, N.S. 1965. Further records of the Four-toed Salamander with remarks on its Habitat in Quebec Prov- ince. Canadian Field-Naturalist 79 : 76-77. Dundee, H. A. 1968. First record of the Four-toed Sala- mander, Hemidactylium scutatum, in Mississippi, with comments on its disjunct distribution in Arkansas and Louisiana. Journal of Herpetology 1: 101-103. Gordon, D. M. 1979. New localities for the Northern Spring Salamander and the Four-toed Salamander in Southwestern Quebec. Canadian Field-Naturalist 93(2) : 193-195. Petranka, J. W. 1998. Salamanders of the United States and Canada. Smithsonian Institution Press, Washington and London. 587 p. Sharbel, T. F. 1990. A range extension of the Four-toed Salamander, Hemidactylium scutatum, in Southern Quebec. Canadian Field-Naturalist 105: 285-286. Received 4 October 2000 Accepted 3 June 2002 2002 NOTES 319 Reproductive Characteristics of Northern Pocket Gophers, Thomomys talpoides, in Alberta Alfalfa Fields GILBERT PROULX Alpha Wildlife Research & Management Ltd., 229 Lilac Terrace, Sherwood Park, Alberta T3H 1W3 Canada Proulx, Gilbert. 2002. Reproductive characteristics of Northern Pocket Gophers, Thomomys talpoides, in Alberta alfalfa fields. Canadian Field-Naturalist 116(2): 319-321. . The productivity of Northern Pocket Gopher (Thomomys talpoides) females inhabiting eight alfalfa (Medicago spp.) fields in Alberta was investigated in 1994. Breeding began in late March and was virtually complete by the end of May. A total of 131 pregnant females were captured on all study sites. Females had, on average, 6.1 (standard deviation: 1.4) viable embryos. There were 295 females with placental scars. They had, on average, 5.5 (1.4) scars. In some study sites, the mean number of embryos/female differed from the mean number of placental scars. In a comparison of various Northern Pocket Gopher populations, it is recommended to use mean productivity levels obtained with the same technique, i.e., embryo or placental scar counts. Key Words: Alberta, alfalfa, embryo, litter size, Medicago spp., Northern Pocket Gopher, placental scars, reproduction, Thomomys talpoides. The Northern Pocket Gopher (Thomomys tal- poides) is a fossorial rodent inhabiting hay fields of western Canada and the United States. Because of its burying and feeding activities, it is considered to seriously impact the productivity of certain crops, particularly alfalfa (Medicago spp.) (Proulx 1997). Alfalfa is believed to be an ideal food for this fosso- rial rodent because of its high nutrient and moisture content (Reid 1973). In an alfalfa field in Saskatche- wan, Runnells (1988) reported an average of 6.4 embryos/litter for 17 gravid females, and 6.8 placen- tal scars for 30 other females. Although the Northern Pocket Gopher is common in Alberta (Smith 1993), little is known about its reproduction. The objective of this study was to determine the breeding period, and average litter size of Northern Pocket Gophers inhabiting alfalfa fields in Alberta. Methods This study was part of a research program to investigate Northern Pocket Gopher populations, and develop an effective control strategy (see Proulx 1997). Alfalfa fields were chosen in eight rural com- munities in south-central Alberta: Didsbury, Caroline, Red Deer, Lacombe, Rimbey, Ponoka, Gwynne, and Camrose. The Lacombe alfalfa field was approximately 5 years old with abundant dande- lion (Taraxacum spp.) and orchard grass (Dactylis glomerata). The seven other fields were 2—3 year-old fields dominated by alfalfa. Near the centre of these fields, eight study sites (one 3.6 ha and seven 8 ha each) were established (Proulx 1997). From 18 April to 27 May 1994, all pocket gophers inhabiting the study sites were kill-trapped. In Caroline and Didsbury, no trapping occurred after the spring removal; captured animals consisted only of the breeders inhabiting the study sites. In the six other fields, 20-60-m-wide traplines were established around the study sites, and maintained from 27 May to 22 September to capture animals attempting to colonize the now pocket gopher-free study sites. Therefore, in six study sites, the breeders inhabiting the study sites and those living in areas adjacent to these sites, but still living within the same fields, were captured. Sexual maturity of females was ascertained by determining if the pubic symphysis was open (Hisaw 1924, 1925). General characteristics of uteri were noted (not bred, swollen uterus, uterus with foetus or placental scars). The number of young per litter was determined by counting the number of viable embryos or placental scars. Loeb and Schwab (1987) found in Thomomys bottae that the proportion of pregnancies with resorbing embryos increased as embryo size class increased. The mean crown-rump length of viable and resorbed embryos was used to classify pregnancies as early-medium term (< 21 mm), or late- term (21 mm) (Loeb and Schwab 1987). I used an analysis of variance followed by the Tukey test to compare mean number of embryos and placental scars between populations (Zar 1984). Student f-tests were used to compare mean produc- tivity levels between embryo and placental scar counts (Dixon and Massey 1969). Probability values “P” < 0.05 were considered statistically significant. Results Breeding activities In April, 195 (92%) of 211 females had swollen uterine walls or were pregnant. The first postpartum females were captured during the 17—23 April trap- ping week. The majority (65%) of females (mn = 75) captured during the first week of May had given birth. In Didsbury and Rimbey, breeding populations were trapped by the end of April and no births were recorded. With the exception of one pregnant female ee a 320 THE CANADIAN FIELD-NATURALIST Vol. 116 captured in Lacombe in mid-June, all pregnancies were recorded before the end of May. >0.005 Productivity levels based on viable embryo counts A total of 132 pregnant females were captured from all study sites. Only one pregnant female was captured in Didsbury, and it was not included in comparisons between populations. Females had an ee average of 6.1 (standard deviation: 1.4) viable embryos. However, mean productivity levels dif- fered significantly among study sites (F, ,,,= 4.152, Comparison embryos VS. scars 1692; 0:05 3.806 <0.005 2.266 <0.05 2.458 <0.01 0.174 Range 4—10 1-8 8 8 a ee 6,124 Eke P <0.005) (Table 1). Camrose, Caroline, Ponoka and Gwynne females had the highest mean produc- a tivity. Rimbey, Red Deer and Lacombe had lower ano mean productivity levels that differed only from that ! of Camrose (Table 1). ErS tm ere Only 9 (7%) of 131 females had resorbed embryos, and these were all found in highly produc- tive populations (Table 1). Twelve embryos had been resorbed, and the majority (75%) of them were > 21 mm in length (Table 1). Number of scars SD 1.4 1.4 5.28 Length (mm)t SZ 4 3 a) = Dal 1 ) Productivity levels based on placental scar counts Placental scars were found in 295 females. Mean litter size was 5.5 (+ 1.4) young/female. A signifi- cant difference in mean litter sizes among study sites (Fy 599 = 11.747, P < 0.005) was due to the markedly greater productivity of Camrose females (Table 1). Females from Ponoka, Lacombe, Gwynne and Red Number Number of resorbed embryos of embryos 4 1 5 2 5 2 Deer had similar mean numbers of placental scars Oo 8 E = aa tt = (Table Ly. a 2s Comparison of embryo to placental scar counts The mean number of embryos for 131 females © was significantly (t= 4.435, P< 0.05) greater than lie Dike - eee the mean number of placental scars for 295 females. Sj—- 4+ Ss piles ills oles 1 Pp yes There was no significant difference (P > 0.05) 5 between the mean number of embryos and placental Bel hE ee Oe ee ite scars for Camrose and Red Deer females (Table 1). a However, in females from Gwynne, Lacombe and S (ees TE oie: OE Ponoka, the mean number of scars was significantly By th = Eas as lower than the mean number of embryos (Table 1). o Oo = B & Discussion A E Bh ee a Se Using a gestation period of 18-19 days (Schramm , os 1961; Reid 1973; Andersen 1978), capture data sug- gest that, in 1994, breeding began, at the earliest, & Ss & ot Aree) coment bale near the end of March. Nearly all females captured 5 = § Bi soya ss after mid-May had given birth, and breeding activi- ties in alfalfa fields was virtually completed before June. In Saskatchewan, Runnells (1988) reported similar findings. There, most females were pregnant in April-May, started lactating in May, and finished breeding in June. A slightly longer breeding season . was reported in Colorado, from 1 March to 15 June (Hansen 1960; Reid 1973). The length of the pocket tl Ch) it Gy Cr bred females 0 3 Non- TABLE |. Litter size in northern pocket gopher populations of Alberta alfalfa fields as determined by embryo and uterine scar counts. *Column means with the same letter do not differ (P > 0.05) using Tukey test. +Crown-rump caliper length. o es 5) : : P Biss fs 3 €s a 3 gopher breeding season likely varies among regions BIER SS 8 = ZE according to length of growing season, climate, and a. OO0ANAMe other extrinsic factors (Dixon 1929; Miller 1946; BIA NMtTNOM Desy and Druecker 1979; Runnells 1988). 2002 NOTES Ey! Females of all populations had one litter with an average ranging from 5 to 7 young, depending on embryo or placental scar counts. However, indepen- dent of the technique used, the mean productivity level of Camrose females was consistently the high- est. The higher productivity of these Camrose females may be related to differences in field charac- teristics (soil, moisture) and forage quality, or the age structure of adult female populations (Desy and Druecker 1979; Andersen 1980; Runnells 1988). Female Northern Pocket Gophers inhabiting Alberta alfalfa fields had mean litter sizes that were similar to those of other alfalfa field populations (Hansen 1960; Runnells 1988). These productivity levels were markedly higher than mean litter sizes of 3-5 young reported for native rangelands in Colorado (Hansen 1960; Hansen and Ward 1966). Like Loeb and Schwab (1987), this study showed that embryo resorption occurred in female Northern Pocket Gophers, particularly in litters with large embryos. Embryo resorption could explain differ- ences observed between mean embryo and placental scar counts. Miller (1946) suggested that embryo counts could be overestimated if some dead fetuses were in a preliminary resorption stage and recorded as normal. However, on the basis of Conaway’s (1955) study with laboratory rats, Loeb and Schwab (1987) suggest that resorbed embryos could leave placental scars. Although this study was not designed to explain discrepancies between produc- tivity levels estimated with embryo and placental scar counts, it points out that comparisons between populations should involve mean productivity levels obtained with the same technique. Acknowledgments I thank Pam Cole, Joel Nicholson, Nicole Proulx, Lori Lounsbury, Orest Litwin, Maurice Moore, Paul King, Gordon Walker, Larry Rice, and Kim Nielsen for technical support. Pauline Feldstein and two anonymous reviewers provided helpful comments on an earlier version of the manuscript. Literature Cited Andersen, D. C. 1978. Observations on reproduction, growth and behavior of the northern pocket gopher (Thomomys talpoides). Journal of Mammalogy 59: 418-422. Conaway, C.H. 1955. Embryo resorption and placental scar formation in the rat. Journal of Mammalogy 36: 516-532. Desy, E. A., and J. D. Druecker. 1979. The estrous cycle of the plains pocket gopher, Geomys bursarius, in the laboratory. Journal of Mammalogy 60: 235-236. Dixon, J.S. 1929. The breeding season of the pocket gopher in California. Journal of Mammalogy 10: 327-328. Dixon, W. J., and F. J. Massey, Jr. 1969. Introduction to Statistical analysis. McGraw-Hill Book Company, New York. 638 pages. Hansen, R. M. 1960. Age and reproductive characteris- tics of mountain pocket gophers in Colorado. Journal of Mammalogy 41: 323-335. Hansen, R. M., and A. L. Ward. 1966. Some relations of pocket gophers to rangelands on Grand Mesa, Colorado. Colorado Agricultural Experimental Station Technical Bulletin Number 88. 22 pages. Hisaw, F. L. 1924. The absorption of the pubic symphysis of the pocket gopher, Geomys bursarius (Shaw): American Naturalist 58 (654): 93-96. Hisaw, F.L. 1925. The influence of the ovary on the resorption of the pubic bones of the pocket gopher, Geo- mys bursarius (Shaw). Journal of Experimental Zoology 42: 411-433. Loeb, S. C. 1990. Reproduction and population structure of pocket gophers (Thomomys bottae) from irrigated alfalfa fields. Proceedings Vertebrate Pest Conference 14: 76-81. Loeb, S. C., and R. G. Schwab. 1987. Estimation of litter size in small mammals: bias due to chronology of embryo resorption. Journal of Mammalogy 68: 671-675. Miller, M. A. 1946. Reproductive rates and cycles in the pocket gopher. Journal of Mammalogy 27: 335-338. Proulx, G. 1997. A northern pocket gopher (Thomomys talpoides) border control strategy: promising approach. Crop Protection 16: 279-284. Reid, V. H. 1973. Population biology of the northern pocket gopher. Pages 21-41 in Pocket gophers and Colorado mountain rangeland. Edited by G. T. Turner, R. M: Hansen, V.H. Reid, H. P. Tietjen, and A. L. Ward. Colorado State University, Fort Collins, Bulletin 554S. Runnells, J. 1988. Some aspects of population dynamics within a southern Saskatchewan population of northern pocket gophers (Thomomys talpoides). M.Sc. thesis, University of Regina, Saskatchewan. 116 pages. Schramm, P. 1961. Copulation and gestation in the pock- et gopher. Journal of Mammalogy 42: 167-170. Smith, H. C. 1993. Alberta mammals. An atlas and guide. Provincial Museum of Alberta, Edmonton. 239 pages. Zar, J. H. 1984. Biostatistical analysis. Second edition. Prentice-Hall, Inc., Englewood Cliffs, New Jersey. 718 pages. Received 2 November 2000 Accepted 26 March 2002 322 THE CANADIAN FIELD-NATURALIST Vol. 116 Evidence of a Second Litter in Northern Pocket Gophers, Thomomys talpoides GILBERT PROULX and PAMELA J. COLE Alpha Wildlife Research & Management Ltd., 229 Lilac Terrace, Sherwood Park, Alberta T8H 1W3 Canada Proulx, Gilbert, and Pamela J. Cole. 2002. Evidence of a second litter in Northern Pocket Gophers, Thomomys talpoides. Canadian Field-Naturalist 116(2): 322-323. In June 1995, we captured 12 and14 female adult Northern Pocket Gophers (Thomomys talpoides) in an alfalfa (Medicago spp.) and a pea (Lathyrus spp.) field, respectively, in central Alberta. Mean litter size was significantly higher in the alfalfa population (6.7 + 1.4 young) than in the pea population (4.5 + 1.9 young). Females from the alfalfa field produced only one litter. However, two females from the pea field bore a second litter. This is the first confirmed report of two litters pro- duced during a single growing season by female Northern Pocket Gophers in western Canada. Key Words: Alberta, alfalfa, embryo, Lathyrus spp, litter size, Medicago spp., Northern Pocket Gopher, pea, placental scars, reproduction, Thomomys talpoides. In northern regions, the Northern Pocket Gopher (Thomomys talpoides) has only one breeding period (Tryon 1947; Hansen 1960), which generally lasts from March to June (Runnells 1988; Proulx 2002). Females have only one litter, which ranges in size from 3 to 5 young in native rangelands (Hansen 1960) to more than 6 in alfalfa fields (Runnells 1988; Proulx 2002). From 12 to 18 June 1995 during an evaluation of the capture efficiency of various killing traps, we collected several Northern Pocket Gopher adult females in a pea (Lathyrus spp.) crop and an alfalfa (Medicago spp.) field near Vegreville, approximate- ly 100 km east of Edmonton, Alberta. Reproducing females were recognized by the presence of an open pubic symphysis (Hisaw 1924, 1925). They were autopsied in situ, and the number of young per litter was determined by counting the number of embryos or placental scars. The mean crown-rump length of embryos was measured with a dial caliper. Com- parison between mean litter sizes was done with a Student f-test (Dixon and Massey 1969). We captured 12 females in the alfalfa field. They only had one litter ranging from 5 to 10 young, and averaging 6.7 (standard deviation = 1.4) young. We captured 14 females in the pea field. Ten of them had given birth, and the number of placental scars ranged from | to 7, and averaged 4.5 (+ 1.9) young. This average was significantly smaller than that of females captured in the alfalfa field (t = 3.310, P<0.05). The four other females from the pea field were pregnant. Two of them had 5 and 10 embryos, measuring 22 and 16cm, respectively. Two other females bore 7 and 4 embryos, but also had 5 and 1 placental scars, respectively. Since the two females had well-developed nipples and appeared to have lactated, it is not believed that these placental scars were the result of a partial litter resorption. A decline in the number of viable embryos usually occurs in the later stages of gestation, when embryos measure >21 mm (Loeb and Schwab 1987; Proulx 2002). In this case, embryos measured 8—12 mm and were at mid-term (Loeb and Schwab 1987). Finding a second litter in the pea field population was a surprise because Proulx (2002), after examin- ing 426 females in highly productive alfalfa fields, reported only one litter per female. Previous studies have reported only a single litter a year for the Northern Pocket Gopher (Chase et al. 1982) although it has been suggested that the species could produce more than one litter a year in mild climates (Cahalane 1947; Bonar 1995). To our knowledge, this is the first confirmed case of a second litter in Northern Pocket Gophers. The production of two litters by pocket gophers in Alberta is not in agreement with the gen- eral life history of the species. In northern latitudes, most breeding activities are complete in June (Runnells 1988; Proulx 2002), and young do not leave the maternal burrow before mid-summer (Proulx 1997). Pocket gophers conceived in mid- summer would therefore leave the maternal burrow in late summer-early fall, when the quality and quantity of green vegetation are reduced (Proulx 1998). The survival rate of these late dispersers would likely be lower than that of juveniles born earlier in the year. The reasons for a second litter in a pea field popula- tion are unknown. Literature Cited Bonar, R. E. 1995. The Northern Pocket Gopher — most of what you thought you might want to know, but hesi- tated to look up. United States Department of Agri- culture, Forest Service, Technology and Development Report, Missoula, Montana. 62 pages. Cahalane, V.H. 1947. Mammals of North America. MacMillan Company, New York. 682 pages. Chase, J. D., W. E. Howard, and J. T. Roseberry. 1982. Pocket gophers. Pages 239-255 in Wild mammals of North America. Edited by J. A. Chapman and G. A. Feldhamer. The Johns Hopkins University Press, Balti- more, Maryland. 2002 NOTES 325 Dixon, W. J., and F. J. Massey, Jr. 1969. Introduction to statistical analysis. McGraw-Hill Book Company, New York. 638 pages. Hansen, R. M. 1960. Age and reproductive characteris- tics of mountain pocket gophers in Colorado. Journal of Mammalogy 41: 323-335. Hisaw, F. L. 1924. The absorption of the pubic symphysis of the pocket gopher, Geomys bursarius (Shaw). Ameri- can Naturalist 58 (654): 93-96. Hisaw, F. L. 1925. The influence of the ovary on the resorption of the pubic bones of the pocket gopher, Geomys bursarius (Shaw). Journal of Experimental Zoology 42: 411-433. | Loeb, S. C., and R. G. Schwab. 1987. Estimation of litter size in small mammals: bias due to chronology of embryo resorption. Journal of Mammalogy 68: 671-675. Proulx, G. 1997. A northern pocket gopher (Thomomys talpoides) border control strategy: promising approach. Crop Protection 16: 279-284. Proulx, G. 1998. Evaluation of strychnine and zinc phos- phide baits to control northern pocket gophers (Thom- omys talpoides) in alfalfa fields in Alberta, Canada. Crop Protection 17: 135-138. Proulx, G. 2002. Reproductive characteristics of northern pocket gophers, Thomomys talpoides, in Alberta alfalfa fields. Canadian Field-Naturalist 116(2): 319-321. Runnells, J. 1988. Some aspects of population dynamics within a southern Saskatchewan population of northern pocket gophers (Thomomys talpoides). M.Sc. thesis, University of Regina, Saskatchewan. 116 pages. Tryon, C. A., Jr. 1947. The biology of the pocket gopher (Thomomys talpoides) in Montana. Montana Agri- cultural Experimental Station Technical Bulletin 448. 3 pages. Received 21 November 2000 Accepted 26 March 2002 Narwhal, Monodon monoceros, Near Western Baffin Island, Nunavut, Canada MARK L. MALLORY! and ANDREW B. DIDIUK2 'Canadian Wildlife Service, P.O. Box 1870, Iqaluit, Nunavut XOA OHO Canada 2Canadian Wildlife Service, 115 Perimeter Road, Saskatoon, Saskatchewan S7N 0X4 Canada Mallory, Mark L., and Andrew B. Didiuk. 2001. Narwhal, Monodon monoceros, near western Baffin Island, Nunavut, Canada. Canadian Field-Naturalist 116(2): 323-325. In August 2000, a Narwhal, Monodon monoceros, was found dead on the west coast of Baffin Island at a location where this species had not been previously documented. The Narwhal was very dark for an adult, and it had an injury on its back that may be consistent with being speared by either a harpoon or by another Narwhal. Key Words: Baffin Island, Narwhal, Monodon monoceros, range, injury. Knowledge of the abundance and distribution of various wildlife species in arctic Canada is incom- plete, in large part due to the isolation of some regions and the expense incurred in surveying these sites. Hence, opportunistic wildlife observations from varied field projects can be of considerable assistance in filling in the gaps in species range maps. On 8 August 2000 the Canadian Wildlife Service was conducting helicopter surveys of coastal wetland habitats to verify vegetation characteristics identified in LANDSAT 5 satellite images in and near the Dewey Soper Migratory Bird Sanctuary on the west coast of Baffin Island. Four biologists and a pilot used a Bell Long Ranger helicopter equipped with floats, and navigated using GPS. Just north of the mouth of the Koukdjuak River, at 66° 59’ N, 72°49’ W (Figure 1), we observed a dead Narwhal (Monodon monoceros) near the shoreline, and we landed beside it to investigate. We knew the Nar- whal had washed ashore the previous night, because we had flown over this portion of shoreline at low altitude the previous day on the way to a fuel cache, and on the pale, flat, clay shoreline the Narwhal was an obvious feature. The carcass was in excellent condition, suggesting a recent death. Blood was flowing from recent peck- ing at the eyes and blowhole by larid (gull) or corvid (crow) scavengers. Closer inspection revealed two curious characteristics of this male Narwhal. Al- though we had no measuring equipment with us, we extrapolated measurements from digital photos and determined that the Narwhal had a tusk that was approximately 1.4 m long, and a total body length of approximately 3.9 m, indicating it was an adult (Hay and Mansfield 1989). However, adults of this size and tusk length are typically light coloured on their ven- tral surface and dark, mottled gray on their dorsal sur- face (Banfield 1974; Hay and Mansfield 1989). This male had only two small, whitish patches under the 324 Foxe Basin Cape Dorset THE CANADIAN FIELD-NATURALIST Vol. 116 | Legend @ Town * Narwhal location Narwhal distribution eo. Maluit KS Ee \Kimmirut / “Ses Hudson Strait Re ee, FIGURE 1. Known range of Narwhal (Monodon monoceros) near Foxe Basin, and location of our observation. chin (each approximately 10 x 15 cm), and it was otherwise a uniform mottled gray over the rest of the body, which is an odd colouration for an adult (K. Hay, Nunavut Wildlife Management Board, personal communication). Also, there was a puncture wound on the dorsal surface of the Narwhal centered approximately 2 cm to the right of the spine. The puncture was almost a perfect, smooth circle with a diameter of approxi- mately 5 cm, and it was clearly made as something penetrated from the dorsal surface directly into the central organs of the Narwhal. We could see 10 cm into the wound, at which point the mass of internal tissues had collapsed and sealed the hole. It did not appear to result from a bite from another animal, and thus we considered three possible explanations: gun- shot wound, harpoon wound, or puncture from the tusk of another Narwhal. First, all observers agreed that, given the location, shape and direction of the puncture, the pattern was inconsistent with that expected from a gunshot wound. Second, the wound configuration was consistent with a harpoon punc- ture. However, if this was an animal that had suf- fered internal wounds during an Inuit hunt, it would have travelled at least 200 km from the known range of Narwhal and Inuit hunting regions in this condi- tion. Finally, we postulate that the wound could have resulted from a puncture by the tusk of another Narwhal. The function of the Narwhal tusk has been the topic of considerable speculation, but it is likely used as a weapon, as a sexually selected trait that indicates male dominance (Gerson and Hickie 1985), and perhaps for digging for food. One study has reported injury of a female Narwhal from another Narwhal’s tusk (Ford and Ford 1986). Unfortunately, in our case the lack of dissection and preservation equipment and the approach of a nearby Polar Bear (Ursus maritimus) precluded a detailed examination of the wound, and thus we could not discriminate further between our two possible explanations. Aside from the curious physical characteristics of this carcass, the location of the finding was also noteworthy. Range maps and traditional ecological knowledge of Inuit in nearby communities suggest Narwhal inhabit western Foxe Basin (Figure 1; Strong 1988; Hay and Mansfield 1989; Stewart et al. 1995). This is the first report of a Narwhal along the eastern coast of Foxe Basin (P. Richard, Department of Fisheries and Oceans, personal communication). It is unclear whether this sighting represents a chance observation of a carcass within the species’ normal geographic range, or if instead this was an injured individual Narwhal that strayed from its usual range. Shallow waters and extensive tidal flats are considered atypical habitat for Narwhal (Hay and Mansfield 1989), although they do enter these areas to escape Killer Whales, Orcinus orca (Campbell et al. 1988). Nonetheless, with no communities along 2002 NOTES 325 this coastline and minimal Inuit hunting in this area, it is possible that Narwhal visit this area of Foxe Basin without detection. Acknowledgments We thank Ruben Bowles and Grigor Hope for field assistance, Al Fontaine for preparing the map, and Dale Caswell and two referees for manuscript reviews. Keith Hay of the Nunavut Wildlife Man- agement Board, and Department of Fisheries and Oceans scientists Pierre Richard and Patrice Simon provided valuable comments from unpublished data on this species. Thanks also to Carolyn Mallory of Nunavut Arctic College for help in finding Narwhal “gray” literature. This project was supported in part by the Arctic Goose Joint Venture and the Canadian Wildlife Service (Prairie and Northern Region). Literature Cited Banfield, A. W. F. 1974. The Mammals of Canada. Uni- versity of Toronto Press, Toronto, Canada. 438 pages. Campbell, R. R., D. B. Yurick, and N. B. Snow. 1988. Predation on Narwhals, Monodon monoceros, by Killer Whales, Orcinus orca, in the eastern Canadian Arctic. Canadian Field-Naturalist 102: 689-696. Gerson, H. B., and J. P. Hickie. 1985. Head scarring on male narwhals (Monodon monoceros): evidence for ag- gressive tusk use. Canadian Journal of Zoology 63: 2083-2087. Hay, K.A., and A. W. Mansfield. 1989. Narwhal Mono- don monoceros Linneaus, 1758. Pages 145-176 in Handbook of Marine Mammals, Volume 4 Edited by S. H. Ridgway and R. Harrison. Academic Press, London. Stewart, D.B., A. Akeeagok, R. Amarualik, S. Panipakutsuk, and A. Taqtu. 1995. Local knowledge of beluga and Narwhal from four communities in arctic Canada. Canadian Technical Report of Fisheries and Aquatic Sciences 2065. 48 pages. Strong, J.T. 1988. Status of the Narwhal, Monodon monoceros, in Canada. Canadian Field-Naturalist 102: 391-398. Ford, J., and D. Ford. 1986. Narwhal: unicorn of the arctic seas. National Geographic 169: 354-363. Received 16 February 2001 Accepted 26 April 2001 A Tribute to Edward Warren Greenwood (1918-2002), Canadian Orchidologist Joyce M. REDDOCH and ALLAN H. REDDOCH Reddoch, Joyce M., and Allan H. Reddoch. 2002. A tribute to Edward Warren Greenwood (1918-2002), Canadian orchidol- ogist. Canadian Field-Naturalist 116(3): 326-330. Ed Greenwood made significant contributions to the knowledge of Canadian and Mexican orchids and their habitats, and was a member of The Ottawa Field-Naturalists’ Club for almost 40 years. He died in Ottawa on 24 February 2002, two weeks after his 84th birthday. He is survived by Mary, his wife of almost 60 years. Edward Warren Eugene Greenwood was born in Toronto, Ontario, on 9 February 1918. He grew up in the rural setting of Stratford, 120 km to the west. During his high school years Ed became interested in the local orchids and in photography, interests which motivated him wherever he lived. With the support of a high school teacher, he built his own pin hole camera. His first photograph was of a Fragrant White Orchid (Platanthera dilatata) growing in the fen margin of a kettle lake just east of town. Also while in high school, Ed met Scottish-born Mary Scobbie, a fellow student and his future spouse. He gave Mary a Showy Lady’s-slipper (Cypripedium reginae) flower for her sixteenth birthday. On graduation from high school, Ed won a schol- arship to study engineering chemistry at Queen’s University, Kingston, Ontario. After his first year he worked in industry for two years before returning to his studies. In 1942 he and Mary were married. A year later Ed graduated with a B.Sc. and entered the army. He was sent to Officer Training School and then continued his studies at the Royal Military College of Science in England. After the war he resigned his commission but continued in research and development as a munitions specialist with the Defence Research Board at Valcartier, Quebec, from 1947 to 1959. While at the Valcartier base, he spent his spare time botanizing and photographing his dis- coveries. He located 14 species of wild orchids with- in the base perimeter, including the rare Southern Twayblade (Listera australis). This population and four others that Ed found in the Quebec City area were the subject of his first scientific paper. Before that, Southern Twayblade was known from only two locations in Canada. After a posting to the Canadian Embassy in Washington, D.C. from 1959 to 1962, Ed and Mary came to Ottawa. Soon after Ed joined The Ottawa Field- Naturalists’ Club in 1963, he got involved in vari- ous local activities including organizing and leading botanical outings and study groups, contributing to members’ slide nights and serving as guest speaker at the 1966 Annual Dinner. On their own, the Greenwoods ranged through eastern Ontario and western Quebec in their Volkswagen Beetle to find orchids and other interesting plants. In the mid-sixties the Club found itself defending the Mer Bleue peat bog from being turned into a garbage dump and attempting to guide the National Capital Commis- sion to an awareness of the ecological treasures in Gatineau Park and the Greenbelt. Ed realized that it would be useful for conservation purposes as well as for orchidological studies to have a catalogue of the orchid colonies and their locations. And so, in 1966 he founded the Club’s Native Orchid Location Survey to seek out and map accurately the orchids within 50 km of the Peace Tower (the Ottawa District). That year he and six others recorded 913 colonies. Buoyed by the success of his initia- tive, Ed decided to expand the survey work to include the whole of Canada. By the mid 1970s some forty people had generated 3000 records in the Ottawa District and a total of 12 000 for all of Canada. This data has proven valuable in preparing submissions for land use management studies and, in particular, for our monograph on the orchids in the Ottawa District (1997, Canadian Field- Naturalist). We have deposited copies of these records in the Agriculture Canada herbarium (DAO) and the herbarum of the Canadian Museum of Nature (CAN). When Ed Greenwood was the guest speaker at the 1966 Ottawa Field-Naturalists’ Club Annual Dinner, his topic was Mexican Wildflowers. He talked about the flowering plants typical of the various vegetation zones of the country and used slides from his exten- sive slide collection to illustrate his subjects. Since the late 1950s the Greenwoods had spent Ed’s annual leave in Mexico and, being excited by what they found, decided to spend their retirement there. Each year they chose a different part of the country to explore and botanize. Ed expanded on his longtime interest in the cactus family and has a species and a variety named after him. In 1969 the Greenwoods left ‘ Ottawa for London, England, where Ed was attached to the Canadian High Commission. During his four years there he spent most weekends at the Royal 326 2002 TRIBUTE TO GREENWOOD 327 Edward W. Greenwood (left) with Mexican field colleague Octavio Suarez in Oaxaca, November 1980. Photograph by Ralph T. Holman. Botanic Gardens, Kew and the British Museum (Natural History) photocopying all available original literature on Mexican orchids and photographing type specimens. Thus equipped, the Greenwoods moved to Mexico when Ed retired in 1973. From their base in Oaxaca, Ed and Mary toured far into the surrounding countryside searching for orchids and intriguing habitats. They also hosted and guided many visiting scientists to areas of interest. Ed was an important contributer to the renaissance of Mexican orchidology begun by Eric Hagsater, who revived the Asociacion Mexicana de Orquideologia and its journal, Orquidea, as well as founding a herbarium (AMO). By 1979 Ed was involved in the journal as book review editor and, with Mary, as translator (to English) and unofficial assistant editor. The journal continued to evolve into a fully refereed, mainly taxonomic publication; Ed served as co-edi- tor for Volumes 8 to 16, from 1981 to 2002. Among Mexico’s thousand or so orchid species, the terrestri- al orchids were the least studied and so Ed focussed his energies on these, especially the genus Govenia, but also the genus Malaxis and the subtribe Spiranthinae. In all, he described 10 new species and clarified the identities of several others. He wrote the account of Govenia for the Flora of North America and advised FNA editor George Argus on various aspects of tropical orchids. After 20 stimulating and successful years in Mexico, Ed and Mary were forced by declining health to return to Ottawa in 1993. There Ed continued to write and to keep in contact with his many friends and colleagues. His remaining papers, two of which are listed in the following Appendix, will be published posthumously. Ed’s Mexican colleagues recognize him as the most influential person in contemporary Mexican orchidol- ogy. They have already named nine orchid species and one genus after him. The next number of /cones eae eee eae cree coe —™” 328 Orchidacearum (Fasciles V and VI, Orchids of Mexico, Parts 2 and 3) will be dedicated to him and will include additional species named in his honour, as well as the two articles he had prepared in Icones format (Eric Hagsater, personal communication). Mexico was Ed’s spiritual home. Ed had a commanding personality, which may be evident in the photograph in Plate 23 of Luer’s The Native Orchids of the United States and Canada (1975). He had a good sense of humour and was a great story teller. He will be remembered by orchid taxonomists for his advocacy of working with living plants in addition to herbarium material and his exhortation to pay special attention to the morpholo- gy of orchid’s reproductive organ, the column. He was insistent that the description be comprehensive and include a detailed technical drawing showing all parts of the plant including several views of the col- umn. Through his enthusiastic promotion of these aspects of orchid study and his eagerness to share information, he had a significant and very beneficial influence on orchid research. His influence as a reviewer and correspondent was remarkable. His energy, enthusiasm, encourgement and expertise, as well as his passion for orchids, will be remembered by many who knew him. Acknowledgments We based this account on Ed’s publications and his voluminous correspondence with us since 1972; on our memories of our friendship with him from 1967 onward; and on information provided by Mary Greenwood, George Argus, Paul Catling, Bill Gamblin, Eric Hagsater, Henry Howden, Dave Smythe, and Marilyn Light, as well as Marilyn’s notes on a 1998 interview with Ed. The photograph came from Eric Hagsater by way of Marilyn Light and Michael MacConaill. Bibliography of E. W. Greenwood [Pagination for Orquidea (Mexico), Volumes 7-11, includes both Spanish and English versions]. Scientific Papers and Notes Greenwood, E. W. 1962. Occurences of the orchid Listera australis in the vicinity of Quebec City. Canadian Field- Naturalist 76(4): 199-202. Greenwood, E. W. 1964. Notes on the phototropic orien- tation of the pseudocephalium of two Mexican species of Cephalocereus. Cactaceas y Suculentas Mexicanas 9(1): 3-6. [in Spanish]. Greenwood, E. W., and H. Sanchez-Mejorada. 1965. Vegetation zones north of Barranca de Metztitlan, Hidalgo, Mexico. Cactaceas y Suculentas Mexicanas 10(4): 92-98. [in Spanish]. Greenwood, E. W. 1967. Orchid Location Survey. Trail & Landscape 1(1): 26-27. Greenwood, E. W. 1967. Plant conservation in the Ottawa Greenbelt and Gatineau Park. Trail & Landscape 1(3): 70-72. THE CANADIAN FIELD-NATURALIST. Vol. 116 Greenwood, E. W. 1967. Mass occurrences of the fern, Ophioglossum vulgatum, in the Ottawa District, Ontario. Canadian Field-Naturalist 81(3): 186-188. Greenwood, E. W. 1968. OFNC location survey of native orchids in Canada. Trail & Landscape 2(4): 98. Greenwood, E. W. 1968. Prairie White Fringed-orchid. Trail & Landscape 2(4): 110-111. Greenwood, E. W. 1968. An orchid new to Gatineau Park: Auricled Twayblade, Listera auriculata Wiegand. Trail & Landscape 2(5): 134-136. : 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. Greenwood, E. W. 1974. Broad-leaved Helleborine now present in Manitoulin District, Ontario. Canadian Field- Naturalist 88(1): 87-88. Greenwood, E. W. 1974. Orchis rotundifolia: addition to the list of plants of the Bruce Peninsula. Canadian Field- Naturalist 88(1): 90. Greenwood, E. W., and R. Gonzalez Tamayo. 1978. Malaxis pollardii L.O. Williams. Orquidea (Mexico) 7(1): 42-51. Greenwood, E. W. 1981. Govenia in Mexico, an introduc- tory note. Orquidea (Mexico) 8(1): 107—120. Greenwood, E. W., and R. Gonzalez Tamayo. 1981. Malaxis amplexicolumna Greenwood & Gonzalez; a new species from Chiapas, Mexico. Orquidea (Mexico) 8(1): 121-136. Balogh, P. Burns, and E. Greenwood. 1982. Cutsis Balogh, Greenwood, and Gonzalez, a new genus from Mexico. Phytologia 51: 297-298. Greenwood, E. W. 1982. Viscidium types in the Spiranthinae. Orquidea (Mexico) 8(2): 283-310. Greenwood, E. W. 1982. Liparis draculoides Greenwood: a new species from Oaxaca, Mexico. Orquidea (Mexico) 8(2): 311-326. Greenwood, E. W. 1983. Artorima erubescens (Lindl.) Dressler & Pollard notes on the pollination mechanism. Orquidea (Mexico) 9(1): 113-122. Greenwood, E. W., and R. Gonzalez Tamayo. 1983. Malaxis javesiae (Reichb. f.) Ames. Orquidea (Mexico) 9(1): 123-142. Greenwood, E. W. 1983. Mexican terrestrial orchids. Pages 81-86 and 44-45 (photographs) in North American terrestrial orchids. Edited by E. H. Plaxton. Michigan Orchid Society, Livonia. Gonzalez Tomayo, R. and E. W. Greenwood. 1984. Malaxis rosilloi, a new species from western Mexico. Orquidea (Mexico) 9(2): 387-395. Greenwood, E. W. 1986. Ponthieva angustipetala Greenwood, an unexpected new species from southern Mexico. Orquidea (Mexico) 10(1): 7—26. Greenwood, E. W. 1987. Govenia bella, a new species from Oaxaca, Mexico. Orquidea (Mexico) 10(2): 229-246. Greenwood, E. W. 1987. Habenaria ixtlanensis, a new species from southern Mexico. Orquidea (Mexico) 10(2): 297-316. Catling, P. M., and E. W. Greenwood. 1988. Wasps and flies in orchid capsules. Orquidea (Mexico) 11: 277-279. Soto Arenas, M. A., and E. W. Greenwood. 1989. Undesirable technical terminology - a current example. Orchid Research Newsletter 13: 8-9. Greenwood, E. W. 1990. Pelexia scintillans, a new species from Oaxaca. Orquidea (Mexico) 12(1): 47-53. 2002 TRIBUTE TO GREENWOOD 329 Greenwood, E. W. 1990. Cranichis schaffneri: \ectotypi- fication and transfer to Ponthieva. Orquidea (Mexico) 12(1): 54-56. Greenwood, E. W. 1990. Govenia bella in Icones orchi- dacearum Fascicle 1. Orchids of Mexico Part 1. Edited by E. Hagsater and G. A. Salazar. Asociacion Mexicana de Orquideologia A.C. Greenwood, E. W. 1991. The Florida Govenia. American Orchid Society Bulletin 60(9): 867-869. Greenwood, E. W. 1992. Govenia liliacea: description and neotypification of a much confused species. Orquidea (Mexico) 12(2): 155-168. Greenwood, E. W. 1992. Govenia capitata Lindley, a central Mexican endemic. Orquidea (Mexico) 12(2): 169-177. Greenwood, E. W. 1992. Malaxis urbana, a new species from Oaxaca. Orquidea (Mexico) 12(2): 199-204. Salazar, G. A., and E. W. Greenwood. 1993. Govenia praecox, a new orchid species from Veracruz. Orquidea (Mexico) 13(1—2): 113-120. Greenwood, E. W. 1993. Govenia dressleriana, another new species rescued from confusion. Orquidea (Mexico) 13(1-2): 165-172. Greenwood, E. W. 1996. The Florida Govenia. North American Native Orchid Journal 2(4): 344-349. Greenwood, E. W. 2002. Govenia. Pages 638-639 in Flora of North America north of Mexico. Volume 26. Edited by Flora of North America Editorial Committee. Oxford University Press, New York. Greenwood, E. W., M. A. Soto, and G. A. Salazar. [in press] The genus Euchile neotypification of Sobralia cit- rina and description of two new species. Icones Orchi- dacearum. Greenwood, E. W. [in preparation] Malaxis plantagoides, an unusual new species from Central Oaxaca, Mexico. Scientific Documents [Internet] Greenwood, E. W. 2000. Tsochilus.http://www.geocities.com/brassia.geo/892000. html. Note, photographs and drawings. Tributes and Book Reviews Greenwood, E. W. 1978. Glenn E. Pollard, student of Mexican orchids. Orquidea (Mexico) 7(1): 3-10. Greenwood, E. W. 1979. Glenn E. Pollard - student of Mexican orchids. American Orchid Society Bulletin 48(1): 49-51. Greenwood, E. W. 1980. Anne Hanes, first editor of T&L. Trail & Landscape 14(1): 2. Greenwood, E. W. 1981. Guido Federico Joao Pabst, orchidologist. Orquidea (Mexico) 8(1): 93-96. Greenwood, E. W. 1992. Dr. Louis O. Williams. Orquidea (Mexico) 12(2): 127-129. Greenwood, E. W. 1964. [Book review] Cacti and other succulents by R. Ginns. Canadian Field-Naturalist 78(2): 121. Greenwood, E. W. 1965. [Book review] Orchids of the Western Great Lakes Region by F. W. Case Jr. Canadian Field-Naturalist 79(2): 149-150. Greenwood, E. W. 1979. [Book review] Native orchids of North America north of Mexico by D. S. Correll. Orquidea (Mexico) 7(3): 262-264. Greenwood, E. W. 1981. [Book review] Notes on the Brachypetalum group of Paphiopedilum by D. C. Erdmann. Orquidea (Mexico) 8(1): 104-106. Greenwood, E. W. 1981. [Book review] Indian orchids: guide to identification and culture by U. C. Pradhan. Orquidea (Mexico) 8(1): 137-139. Greenwood, E. W. 1981. [Book review] Orchids of India by T. K. Bose and S. K. Bhattacharjee. Orquidea (Mexico) 8(1): 140-142. Greenwood, E. W. 1982. [Book review] The orchids, nat- ural history and classification by R. L. Dressler. Orquidea (Mexico) 8(2): 399-405. Greenwood, E. W. 1983. [Book review] Orchid biology: reviews and perspectives, II. Edited by J. Arditti. Orquidea (Mexico) 9(1): 165-174. Greenwood, E. W. 1983. [Book review] Southern African epiphytic orchids by J. S. Ball. Orquidea (Mexico) 9(1): 175-178. Greenwood, E. W. 1983. [Book review] A revised hand- book to the flora of Ceylon, Volume 2. Edited by M. D. Dassanayake and F. R. Fosberg. Orquidea (Mexico) 9(1): 179-184. Greenwood, E. W. 1983. [Book review] Orchids from Curtis’s Botanical Magazine by D. R. Hunt. Orquidea (Mexico) 9(1): 185-190. Greenwood, E. W. 1984. [Book review] Miniature orchids by R. T. Northen. Orquidea (Mexico) 9(2): 346, 396. Greenwood, E. W. 1986. [Book review] Wild orchids of southern Africa by J. Stewart, H. P. Linder, E. A. Schelpe and A. V. Hall. Orquidea (Mexico) 10(1): 156-160. Greenwood, E. W. 1986. [Book review] Orchidaceae: Volume 16, Flora Novo-Galiciana by R. McVaugh. Orquidea (Mexico) 10(1): 206-212. Genus and Species Described by E. W. Greenwood [Compiled by ER1c HAGSATER] ORCHIDACEAE Genus Cutsis. P. Balogh, E. Greenwood & Gonzales, in Phytologia, 51(5): 297 [1982]. Species Euchile depatronii Greenwood, Soto & Salazar [in press Icones Orchidacearum] Euchile hagsateri Greenwood, Soto & Salazar [in press Icones Orchidacearum] Govenia bella E. W. Greenwood, in Orquidea (Mexico) 10(2): 230 [1987]. Govenia dressleriana E. W. Greenwood, in Orquidea (Mexico) 13(1—2): 165 [1993]. Govenia praecox G. A. Salazar & E. W. Greenwood, in Orquidea (Mexico) 13(1+2): 113 [1993]. Habenaria ixtlanensis E. W. Greenwood, in Orquidea (Mexico) 10(2): 297 [1987]. Liparis draculoides E. W. Greenwood, in Orquidea (Mexico) 8(2): 312 [1982]. Malaxis amplexicolumna E. W. Greenwood & R. Gonzalez Tamayo, in Orquidea (Mexico) 8(1): 123 [1981]. Malaxis rosilloi R. Gonzalez Tamayo & E. W. Greenwood, in Orquidea (Mexico) 9(2): 387 [1984]. Malaxis urbana E. W. Greenwood, in Orquidea (Mexico) 12(2): 200 [1992]. Pelexia scintillans E. W. Greenwood, in Orquidea (Mexico) 12(1): 48 [1990]. Ponthieva angustipetala E. W. Greenwood, in Orquidea (Mexico) 10(1): 9 [1986]. 330 New Combinations by E. W. Greenwood Cyclopogon comosus (Rchb. f.) P. Burns-Balogh & Green- wood, in Orquidea (Mexico) 10(1): 92 [1986]: Notes: Spiranthes comosa. Ponthieva schaffneri (Rchb. f.) E.W. Greenwood, in Orquidea (Mexico) 12(1): 55 [1990]: Notes: Cranichis schaffneri. Schiedeella diaphana (Lindl.) P. Burns-Balogh & Green- wood, in Orquidea (Mexico) 10(1): 93 [1986]: Notes: Spiranthes diaphana. Stenorrhynchos petensis (L.O. Wms.) P. Burns-Balogh & Greenwood, in Orquidea (Mexico) 10(1): 93 [1986]: Notes: Spiranthes petensis. Species Named After E. W. Greenwood CACTACEAE Coryphantha greenwoodii H. Bravo, in Cactaceas y Suculentas Mexicanas xv 27 [1970] Ferocactus recurvus var. greenwoodii, in C. Glass. Cactus and Succulent Journal (US) 40: 160 [1968]. ORCHIDACEAE Greenwoodia sawyeri (Standl. & L. O. Williams) P. Burns- Balogh, in Orquidea (Mexico) 10(1): 80 [1986]. THE CANADIAN FIELD-NATURALIST. Vol. 116 Bletia greenwoodiana V. Sosa, in Brittonia 46(3): 208 (1994). Encyclia greenwoodiana J. Aguirre-Olavarrieta, in Orquidea (Mexico) 12(2): 205 [1992]. Epidendrum greenwoodii E. Hagsater, in Orquidea (Mexico) 10(2): 338 [1987]. Galeandra greenwoodiana N. Warford, in Lindleyana 9(1): 39 [1994]. Habenaria greenwoodiana R.Gonzalez Tamayo, in Boletin del Instituto de Botanica, Universitad Guadalajara 6(2—3): 204 [1998, published 2000]. Lepanthes greenwoodii G. A. Salazar Chavez & M. A. Soto Arenas, in Orquidea (Mexico) 14: 103 [1996] Liparis greenwoodiana A. Espejo Serna, in Orquidea (Mexico) 10(2): 370 [1987] Malaxis greenwoodiana G. A. Salazar Chavez & M. A. Soto Arenas, in Acta Botanica Mexicana 10: 46 [1990]. Trichosalpinx greenwoodiana M. A. Soto Arenas, in Orquidea (Mexico) 10(2): 257 [1987] Additional tribute Light, M. H. S., J. M. Reddoch, E. Hagsater, and M. Soto. 2003. Edward W. Greenwood (1918-2002). Icones Orchidacearum, Fascicles V and VI. Orchids of Mexico, Part 2 and 3. Pages ii—vil. Book Reviews ZOOLOGY Butterflies of British Columbia (including Western Alberta, Southern Yukon, the Alaskan Panhandle, Washington, Northern Oregon, Northern Idaho, Northwestern Montana) By C.S. Guppy, and J. H. Shepard. 2001. Royal British Columbia Museum book published by UBC Press, Vancouver. 414 pp., illus. $95. British Columbia has the largest and most diverse butterfly fauna of any Canadian province with 187 species recorded to date. Chris Guppy and Jon Shepard take on the challenge of detailing all the available information on these charismatic species in their well-researched and attractive book. In the last thirty years, books have been published on most province’s butterflies. British Columbia is one of the last to produce a comprehensive butterfly book, but in many respects it may be the best. The closest in quality is Alberta Butterflies, published in 1995. The 70 pages of introductory chapters cover such standard topics as the study, conservation, gardening and biology of British Columbia butterflies. They also cover less standard topics such as post-glacial origin, impact of humans, the morphology of imma- tures and adults, and seasonal changes of the British Columbia butterfly fauna. Where this book excels is in the copious, useful detail. Each chapter is sub- divided into appropriate headings to make the infor- mation you may be looking for easy to find. I partic- ularly liked the list of butterflies of concern in British Columbia and the series of four maps show- ing how the glacial icecaps covered the province. The species accounts make up the majority of the text. Each covers the etymology ( the origin and meaning of the common and Latin names), immature stages, biology, subspecies, range and habitat, gener- al distribution, and conservation status. The book is arranged with two columns of text for each page. When you first open the book, it is the visuals for each species that attracts your attention. In addition to half-column size photos of the upper and under- sides of the wings for all species, there are full-col- umn excellent pictures from nature of adults and lar- vae for many species. Each account also has a map of British Columbia and surrounding regions show- ing detailed dot locations for all subspecies (a unique feature), differentiated by colour. As well, there is a bar graph showing the relative abundance of each species throughout the known flight season. The final part of the book contains appendices with a species checklist, data and credits for the but- terfly photographs and drawings, the Lepidopterists’ Society statement for collecting lepidoptera, a glos- sary, an extensive bibliography and an index. There are also maps showing the distribution of 20 addi- tional species in areas adjacent to British Columbia that could show up in the province. This additional species concept is slightly confusing because there are also full species descriptions in the text for at least seven species that have not yet been found in British Columbia. As with all such taxonomic books, there are some contentious points. The authors have reassessed the status of several species that differs from the taxono- my of most recent authors. As an example, they treat the population of the Brown Elfin, /ncisalia augusti- nus, in southern British Columbia as a separate species, the Western Elfin, /ncisalia iroides. Also contentious is the use by the authors of common names that have never been used before in the but- terfly literature (e.g., Boeber’s Fritillary and the Mountain Alpine). Overall, this is an admirable work abounding in useful information and should be on the bookshelves of anyone interested in butterflies and their role in nature. PETER HALL 24 Wendover Avenue, Ottawa, Ontario KIS 4Z7 Canada Ecosystem Dynamics of the Boreal Forest: The Kluane Project Charles J. Krebs, Stan Boutin, and Rudy Boonstra. 2001. Oxford University Press, Oxford. 511 pp., illus. U.S. $152. The 10-year-cycle of the Snowshoe Hare and Lynx has perplexed and intrigued Canadians for nearly 200 years. Why is it so regular, so predictable? WwW Yukon’s Kluane National Park was chosen for a decade of intensive study of the 10-year cycle, “the main dynamic driving the population trends of larger vertebrate predators,” funded mainly by the Natural Sciences and Research Council of Canada. Nine fac- ulty members from three Canadian universities, 26 se graduate students, 75 summer assistants, and 18 tech- nicians expended 158 person-years of effort. There were 98 058 field hours of observations by 212 observers and 2780 small mammals were trapped dur- ing 24,650 trap nights. Tracks were counted along 12 194 km of transect. All the significant parts of the system, including vegetation, were described and quantified precisely in the same place at the same time. Experimental manipulation was used to test hypotheses. Do predators control hare numbers, which in turn regulate the vegetation, known as top- down effects? Or do soil nutrients regulate vegeta- tion, which regulates numbers of hares and thereby the numbers of their predators, so-called bottom-up effects? Or are the effects reciprocal? Every component of the boreal forest community was investigated; five chapters deal mainly with veg- etation. Special attention was given to the biomass of hares and ground squirrels and of the main predators, Lynx, Coyote, and Great Horned Owl. Experimental manipulation of fertilizer, use of herbicide to kill vegetation, food supplementation by addition of rab- bit chow, exclusion of hares, and attempted exclu- sion of carnivorous predators, were studied alone, within enclosures, and in combination. Simulations tested 27 individual models. There was a 26- to 44-fold variation in numbers of hares. Coyotes responded with a 6-fold and Lynx with a 7.5-fold variation. During the decline and low phases of the hare cycle, many predators did not reproduce. As expected, addition of fertilizer caused spruce branches to grow more, and hare densities went up 1.3-fold. Removal of mammalian predators doubled the number of hares, and addition of rabbit chow tripled their number. Chow addition and predator exclusion together caused hare numbers to rise 9.7- fold, advanced hare parturition dates by 5 to 12 days, and increased the number of young weaned per female hare. Food addition postponed the hare popu- lation decline. Most years, over 90% of adult hares that died were killed by predators, and 65 to 75% of these predator killings were by Coyotes and Lynx. When these mammals were fenced out, Great Horned Owls and Northern Goshawks accounted for 70 to 80% of the predator-killed hares. Great Horned Owls were almost entirely dependent on Snowshoe Hares dur- ing winter, killing at least 5 to 10% of hares present in peak hare years, and up to 38% during one year of hare decline (1992-93). Great Horned Owls were competitively superior to Lynx and Coyotes. THE CANADIAN FIELD-NATURALIST Vol. 116 There were unexpected findings. Great Horned Owl numbers were limited by food even at the peak of the hare cycle; the owl peak lagged two years after the hare peak. The Northern Goshawk was the only predator that switched from hares to grouse dur- ing the hare decline. Unexpectedly, Arctic Ground Squirrels numbers were in synchrony with the 10- year cycle of the hares. Spruce Grouse peaked a year ahead of the hares. Hare browsing stimulated pro- ductivity of shrubs, rather than diminishing this food supply. Contrary to expectations, sufficient food was available to hares throughout the cycle. It was expected that ground squirrels and red squirrels would function mainly as alternate prey for preda- tors; instead, to everyone’s astonishment, both squir- rel species ate little hares, killing up to three-quarters of them at the peak of the hare cycle. These squirrel kills thus helped to prevent hares from overgrazing their food supply. The expected four-year vole cycle, evident in Europe, was not seen at Kluane; Red- backed Voles had major peaks 11 years apart, inversely related to hare populations. This study provides an almost overwhelming wealth of new information. It offers the first-ever evidence of Lynx and Coyotes acting together as a predator guild, accounting for 78% of mortality of radio-collared hares over 11 years. Coyote numbers were highest at the peak of the hare cycle, but Lynx lagged a year behind. Predators appeared to induce intense stress in hares, thereby depressing reproduc- tion to a degree not previously recognised. The authors conclude that the boreal forest com- munity is predominately a top-down system. The major changes are driven by predators; predators could not survive in the boreal forest in the absence of hare peaks. Most hares die from predation and very few from starvation. The reasons the cycle averages ten years between peaks still have not been explained, except for a tantalizing paragraph that suggests “intrinsic” programming of hares! Data and computer simulations are supplied on a CD-ROM, supplied with each book purchased. This book, which provides extremely detailed information about one of biology’s greatest puzzles, is an absolute must for all University libraries, in spite of its almost prohibitive price. Those for whom the ten-year cycle has held a lifelong fascination will wish at least to borrow it. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N 0J8 ’ Canada 2002 BOOK REVIEWS 333 The Canadian Rockies Guide to Wildlife Watching: The Best Places to See and Appreciate Animals in Their Natural Habitat By Michael Kerr. 2000. Fifth House Publishers, Calgary. 237 pp., illus. $26.95. Kerr, a wildlife biologist with the Canadian Forest Service, communicates his love of the Rocky Mountains in a book that is partly personal reflec- tion, partly wildlife biology, partly a travelogue, and most of all a description of the opportunities and eti- quette of wildlife observation in the Canadian Rockies. The book has large print, good pictures and maps, and detailed information about roads, towns, munici- palities, and parks. It is divided into four sections, the first of which introduces the idea of the book and why it needs to be written at this time. The second section provides a general description of a large number of fish, reptiles, birds, and larger animals resident to the Canadian Rockies. The third section outlines areas where wildlife may be viewed. Finally there is a calendar showing viewing opportunities in each month of the year, describing migration pat- terns, and identifying prime viewing locations. Even though a small book on a distinct region, it contains much detail and answers many questions on watch- ing wildlife in the Rocky Mountains. The book was a fairly quick read suited to an arm- chair on a quiet evening or as a ready reference on a road-trip, with an interesting presentation style (sometimes tongue-in-cheek) which kept the reading interesting and made me want to go back and read some parts again. As a wildlife and bird enthusiast, I was more interested in his comments about viewing and encountering large mammals and different bird species. As a good biologist and honest reporter, Kerr was frank about the possibilities of seeing mountain goats and large cats up close, or even see- ing them at all. This book is not about a safari where all animals are promised and delivered. Mountain goats are rarely seen up close and often with great difficulty even when using a spotting scope to view mountain ledges. Many wildlife enthusiasts living in the mountains have never in their lives had the good Four Wings and a Prayer By Sue Halpern. 2001. Alfred A. Knopf Canada, Toronto. 224 pp. $29.95. When most people think of butterflies, the first image that usually comes to mind is that charismatic species, the tawny-orange Monarch. The “king” of butterflies continues to fascinate anyone remotely interested in the natural world because of its extraor- dinary migrations. It has attracted some of North America’s best nature writers to chronicle its unique life history. fortune to see any of the big cats while always recog- nising signs of their presence. Only a few weeks after reading the book I encoun- tered a national newspaper article on viewing the migration of Golden Eeagles. From my reading of the book, I was able to identify the area involved in the article and I was also aware of the background of the recent discovery of the birds in migration, and could compare data in the article to the data reported in the book. His discussion of sighting birds and their migra- tions was interesting and like a good tour guide made me think of the possibilities of going to view. Fish and reptile viewing were also included to complete the wildlife inventory. Hunting and fishing seasons were mentioned in passing though this is not a book about capturing wildlife on anything but photographic film. As a further guide to the possibilities of seeing any one of the species named in the book, Kerr has included a “horseshoe” rating to give an idea of the frequency of occurrence of seeing the species. One horseshoe means common sightings, two is less common, three very rare, and four is extremely rare. A wildlife viewer should be prepared with the knowledge of how to identify any wildlife which can be seen but not to expect too much. After any road journey through the mountains, we invariably ask each other “What did you see?” The answer would be governed by opportunity, most often by luck and occasionally include a sighting which was planned or hoped for. Seeing elk in Banff National Park or the townsite is more probable if you look on both sides of the road. This book teaches in a gentle and entertaining way the appreciation of both sides of the road, the slopes, the creeks, and the sea- sons of the year. Kerr would include the respect for animals’ capabilities to defend themselves or to hide from unwanted observers. JIM O’ NEIL St. Mark’s College, 5935 Iona Drive, Vancouver, British Columbia V6T 1J7 Canada This book is the latest in the genre and when I first started to read it, | wondered what new insights could be added about this already well-described phenomenon. Well, Sue Halpern has brought a refreshing new approach. She focusses on the people who interact with the butterfly. The biggest mystery about the Monarch migration is how the species, over at least three generations, finds its way in the fall to a relatively small area in Mexico and then returns as far north as Canada in the 334 spring. As stated on the dust jacket, Sue Halpern “recounts the mesmerizing story of this mystery and the equally appealing and remarkable people who are attempting to solve it”. And she is refreshingly candid about the personalities involved, some of whom I have met and indeed are “characters”. Robert Michael Pyle came out with his own best- seller on the Monarch in 1999 when he published Chasing Monarchs: Migrating with the Butterflies of Passage. Pyle, a white-bearded, elfish “character” from Washington State, took a different approach than Halpern. He got into Marsha, his well-travelled Honda, and started “wandering with the wanderers” from British Columbia to Mexico. Halpern takes her own journeys, joining up with the best known of the Monarch specialists and following them around to where the Monarchs fly in Mexico, Texas, New Jersey, Kansas, and Ontario. Halpern chronicles the relationships among the many Monarch specialists, none more fascinating than that between Fred Urquhart, the doyen and first to discover where the Monarchs overwintered in Mexico, and Lincoln Brower, the distinguished American biologist. Urquhart, from the University of Toronto, had been active for many years tagging the butterflies and attempting to locate the Mexican sites. (As a teenager, I spent a day in fields around Golden Lake in Ontario helping Urquhart stick tags on butterfly wings). Halpern records that Brower was seen by Urghhart as an upstart and wouldn’t shake his hand when they met. Radio Tracking and Animal Populations Edited by J. J. Millspaugh and J. M. Marzluff. 2001. Academic Press, San Diego, California. 474 pp., illus. USS. $69.95. This book updates an earlier Academic Press book Analysis of wildlife radio-tracking data (White and Garrott 1990). It uses the expertise and experi- ence of several scientists who are knowledgeable in radio telemetry and the analysis of telemetry derived data. Among the authors is Dr. G. C. White, the senior author of the earlier book (White and Garrott, 1990). Radio tracking and animal populations covers experimental design, animal movements, resource selection, and population dynamics. A chapter on telemetry equipment provides an overview of the technology who is sufficient for the scientist that is concerned mainly with analysis, but should not supersede another Academic Press volume A manual for wildlife radio tagging (Kenward, 2001). For this book, Dr. R. E. Kenward has provided an excellent chapter on the history of wildlife telemetry. THE CANADIAN FIELD-NATURALIST. Vol. 116 The author also shows how modern technology finds intriguing new ways to expand our knowledge. When not physically with her characters, she tunes into the Internet chat line on Monarchs known as D- Plex (the scientific name for the Monarch is Danaus plexippus) where hundreds of individuals around North America record every sighting of Monarchs they make. Halpern also introduces us to two amaz- ing Canadians. Don Davis has tagged more Monarchs than any single person, an estimated twen- ty thousand, while David Gibo regularly takes his glider plane up into Ontario skies looking for Monarchs to try to better understand their aerody- namics. Halpern takes her fear of flying in hand and goes up with him. Throughout this well-written and thoroughly enjoyable book, Halpern uses humour and her sensi- tivity both to the subject and the use of language to weave a fascinating tale of people interacting with this amazing insect. She chronicles how a myriad of amateur and professional enthusiasts have tried to track down and answer the mysteries surrounding the Monarch. Despite this, she laments that “...physi- cists were going to be able to tell us how the world worked, but we still wouldn’t know how a single Monarch butterfly found its way from Canada to Mexico...” PETER HALL 24 Wendover Avenue, Ottawa, Ontario K1S 4Z7 Canada One of the best features in the very clean and error free volume is the appendix. This appendix provides a comprehensive listing of software avail- able for use in analyzing telemetry data. The listing includes web sites where information on the various software packages can be obtained, and in some cases the software and documentation can be down- loaded. In many cases the appendix provides suffi- cient information to aid in deciding which pro- gram(s) to select for a particular analysis. In addi- tion, the literature cited provides one of the most complete bibliographies of telemetry literature that I have seen in many years. This book is a must reference for all scientists that work with telemetry or are contemplating ‘telemetry projects. It should be used from start to finish in any telemetry project. Coupled with Kenward (2001), you have at your disposal a com- plete telemetry library. Academic Press is to be commended for continuing and updating its wildlife telemetry offerings. 2002 BOOK REVIEWS 335 Literature Cited Kenward, R. E. 2001. A manual for wildlife radio tagging. Academic Press, San Diego, California. White, G. C., and R. A. Garrott. 1990. Analysis of wildlife radio- tracking data. Academic Press, San Diego, California. ROGER D. APPLEGATE Kansas Department of Wildlife and Parks, Research and Survey Office, P.O. Box 1525, Emporia, Kansas 66801 The Birds of Ecuador — Status, Distribution and Taxonomy (Volume I), Field Guide (Volume IT) By Robert Ridgely and Paul Greenfield. 2001. Cornell University Press. xvii + 848 pp. (Volume I) and xvii + 934 pp. (Volume II), illus. Paper U.S. 10$. I was delighted to hear that a new, two volume set on Birds of Ecuador was about to be published because earlier this year I searched for a good field guide to birds of Ecuador, and discovered that this was a neglected area. Les Beletsky (1998) had recent- ly produced a traveler’s guide, but this was more of an ecosystem text than a field guide. True it had a field guide section but this covered only the most common birds (plus reptiles, amphibians, mammals and a handful of key insects). As nice as this book is, it would not fulfill the needs of an avid birder. By default I turned to the Birds of Colombia (Hitty and Brown 1986). This is a very good guide, with excel- lent illustrations. However I began to generate many questions. For example, the Colombia guide contains little or no information on Casqued Oropendula, Purplish Jacamar, and Hauxwell’s Thrush. This new publication has now arrived as a boxed set. The first volume deals with distribution and tax- onomy and the second is called the field guide. So how well does the field guide work? For a start Vorme tl is large; Sicm ‘K 15:.5:cm 22.5 cm (2in X 6in X 9.25 in), weighing over 1.5 kilos (almost 3.5 lbs.). The author confronts this issue head on by explaining he needed the size to include all the 1,600 species of birds. However I would dis- agree for reasons I will discuss later. I believe this volume could be two-thirds to, possibly, half the cur- rent size. The balance of the cut material could go in Volume 1. From a technical point of view, as opposed to the logistics of size, the book fares much better. Finally we have a decent reference to one of the most diverse habitats on earth. Each species has a plumage description of all the salient features and comparisons to similar species. There is a short description of range and a respectably sized, clear range map. There are many species that are not cov- ered or sketchily described in Birds of Columbia that are well covered in these volumes. However these books only cover mainland species. They do not include the Galapagos Islands, although some birds are included, as they are visitors to the mainland coast. There is a colored map of the three main ecore- gions at the beginning of the coloured plates. This map is repeated in black and white in the front of both volumes. I found the location of (San Sebastian de) Coca is not correct (it is shown east of Rio Napo, whereas it is on the west side, on the Rio Coca) but other places seem to be positioned correctly. There is no scale given (the reader can assume the colour bar explaining the elevation code represents about 250 km). The major rivers are shown but not named. You will have to identify the Napo, Aguarico, Zamora, and Guaillabemba, etc. to follow the distri- bution section. These volumes are mostly free of serious technical errors except that the distribution map for Hairy-crested Antbird is repeated. The map for Yellow-billed Tern I think is shaded. It is only possible to see shading with strong magnification. Compare this shading with the easy to view map of the Large-billed Tern, which has a similar distribu- tion. The Black Skimmer is shown only on the Napo whereas the text says it occurs on the Rio Pastaza. A word is in order about the names and status of some specific species. The Great Mannikin is renamed Varzea Schiffornis as the authors suggest and substantiate that this is a more accurate name. The Yellow-ridged Toucan is lumped with the Channel-billed Toucan and the recent combination of the Red-backed and Puna Hawks as the Variable Hawk is also covered. The authors have also changed some of the scientific names to represent current thinking; e.g., Red-throated Caracara is moved from Daptrius to Ibycter and Salvin’s Curassow is moved from Crax to Mutu. The infor- mation on the White-chested Swift is sparse and the author gives a good explanation of the current sight- ings and the potential for a change in status. Several other birds that are poorly known may undergo a split when they have received further study. These include the Russet—backed Oropendula (atrocasta- neus and angustifrons subspecies), the look-alike Subtropical and Scarlet-rumped Caciques (the Red- rumped Cacique is also very similar), the eastern and western forms of the Tropical Gnatcatcher, and two forms of the Yellow-rumped Cacique. I was sur- prised that the author made no mention of a potential split of the White-winged Black Phoebe from the widespread Black Phoebe. Other documents list the 336 Variable Seedeater as present along the Rio Napo. This author has given this form species status as Caqueta’s Seedeater. I have a good reference to a virtually all-black form of this bird and was hoping the Ridgely’s taxonomic section would clarify this dilemma. I still remain somewhat confused. The illustrations are of good quality. The shape and posture of the birds are well captured. I find the colours in my edition over bright. This is not an issue for most species as they are very colourful any- way. How can you make a macaw over bright? It is more of a problem with dull birds, where the shades are more subtle. However, I had no problem instant- ly recognizing a species for what it was meant to be. I have always had problems with other people’s descriptions of birds’ song. This is not the case with this book. I found myself nodding in agreement vir- tually all of the time. For example, Ridgley gives the Ruddy Pigeon’s call as “hit the foul pole.” To my ear this is much closer to the sound than “What’s the matter?” that I have read elsewhere. I was impressed with the author’s description of some sounds I thought impossible to describe (e.g., Paraque). The only exception I could find was Bartlett’s Tinamou, but it was within a typical song variation. I do not agree with the author’s view that the field guide volume could not have been smaller. For example, the text for Little Blue Heron in the North American National Geographic guide is only 40% of that in this guide, and is perfectly adequate for field use. I believe the size could be cut in half by cutting out repetition, using cryptic language, and using short forms for oft-repeated word (e.g., A for adult). Indeed for use as a field guide I would have been happy to remove and use just the illustrations with Sylvia Warblers By Hadoram Shirihai, Gabriel Garballo, and Andreas J. Helbig; illustrated by Alan Harris, edited by Guy M. Kirwan and Lars Svensson, with photographic editing and field photography by David Cottridge. 2001. Princeton University Press, Princeton, New Jersey. 576 pp., illus. US$75. This book has been described as one in the series of Helm-Princeton identification guides, and so it is, but here we have a substantial tome devoted to, not a whole family, but to one genus of some 22 species. Yet it is truly an identification guide, with the bulk of the book offering exhaustive detail on the com- plexities of Sylvia plumage variation. The Sylvia warblers are a genus of Palearctic war- bler species. The genus has its centre of abundance in the Mediterranean area, and it includes some of the more colourful and eye-catching members of the Sylviidae; a family that is better noted for its rather nondescript plumages and identification challenges. THE CANADIAN FIELD-NATURALIST Vol. 116 their cryptic text. This 200-page section has most of what I need in the field. To have the illustrations plus the range maps would be even better. I estimate that this would add about 250 pages. The resulting 3cm X 15.5cm X 23.5 cm size is slightly larger than the Geographic guide. The author has the per- fect answer for what to do with the useful material cut from the field guide. Put it all in volume one. I have tried carrying volume two in the field and it is far too heavy to cart around. I left it behind after the first two hour-long walk in the cloud forest. I did not even think of carrying it in the rain forest.- Instead I used it thereafter as a reference each night and I had no difficulty identifying almost all the birds I saw using the plates and text. I did find that there was some confusion among a few of the Ecuadorian birders, but this appeared to relate to the new names brought about by several splits rather than inconsistencies in the book. My paper copy curled badly in the tropical humidity of the Amazon, but returned to normal once back in Canada. This book is a long-awaited, long-needed, impor- tant contribution to ornithological literature. It is a must for anyone working or visiting Ecuador, Amazonia, and South America and it would make a great gift too. Literature Cited Beletsky, Les. 1998. Costa Rica — The Ecotravellers’ Wildlife Guide. Academic Press. Hilty, Steven, and William Brown. 1986. Guide to Birds of Columbia. Princeton University Press. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Gloucester, Ontario K1J 6K5 Canada It also includes such familiar British species as the Blackcap [S. atricapilla], Garden Warbler [S. borin] and Common Whitethroat [. communis]. This is an exceptional book. It has been over a decade in the making, and this depth of scholarship is evident throughout. Innovations, whether in taxon- omy or simply in such matters as the manner in which molt sequences are portrayed, all give the sense of having been carefully considered and well thought-out. A lengthy introduction with a “User’s Guide” carefully outlines the plan of the work. This is fol- lowed by a 20-page section on the characteristics of ‘the genus, dealing first with phylogeny and biogeog- raphy, and then with general biology and ecology. The genus has been the subject of many ground- breaking studies, perhaps most notably on orienta- tion and migration, and its systematics are still the source of debate. This volume does not attempt to 2002 cover the full range of these topics, which are already available in the major handbooks covering the region, but presents an accurate and up-to-date overview. The authors adopt a “modern” version of the Biological Species Concept, recognizing some new species splits, as allospecies belonging to super- species groupings. All these sections are both packed with information and highly readable. The extraordinarily detailed species accounts form the bulk of the book. Each is 20-30 pages in length, and starts with a brief introduction, giving a summa- ry of the species and its systematics. The heading of field identification includes a section on potential confusion species, followed by a detailed “Field characters and plumages” and concluding with a sec- tion on major pitfalls. All this is condensed into a box with an identification summary. Voice is then covered, with sections on “Practical use” and main repertoire, and copious sonograms. Identification in hand included.a discussion on similar species and a figure showing average measurements and ranges. Subspecies taxonomy includes extensive comments on geographic variation and its associated pitfalls. The section on “Moult, Age and Sex” gives detailed descriptions of each of these [using European termi- nology], with a box giving a key to aging and sexing Bats of Papua New Guinea By Frank J. Bonaccorso. 1998. Conservation International Tropical Field Guide Series 2, Washington, DC (dis- tributed by University of Chicago Press, Chicago). 492 pp., illus. U.S. $40. This book is the second in a series of new tropical wildlife field guides published by Conservation International. It is a pleasant surprise that the organ- isms of study are not birds or large mammals, the usual subjects of field guides, but bats! These unique animals are the only mammals that have evolved true powered flight, and a host of associated ecological, behavioural, and anatomical novelties. They are not only interesting from a scientific point of view but also in terms of the conservation of biological diver- sity and the environment. For example, bats are important components of tropical forest regeneration as seed dispersers and flower pollinators. This is most evident in a developing country like Papua New Guinea, the area of focus for this field guide, which has been able to retain most of its natural and cultural wealth in spite of mounting economic pres- sure to unsustainably exploit it. This book aims to promote the development of ecotourism and ulti- mately the preservation of nature. There are 91 species of bats documented from Papua New Guinea, the country that occupies the eastern half of the island of New Guinea, off the BOOK REVIEWS 337 the species, and with an excellent and innovative plate showing the timing and percentages of wing feather replacement at each moult. Finally each account has headings for general biology and ecolo- gy, and population size and trends, and concludes with 2-3 pages of biometrical data. Supplementing this formidable body of text are plates: a figure giving lively vignettes showing the “jizz and movement” of the species; an excellent plate showing plumages of both sexes and juveniles, and different races as appropriate; 3—4 pages of superb photographs — usually 15 to 20 images — and full-page maps showing both breeding and winter ranges. In all, a major tour de force. My only regret is that this is a volume that neces- sarily will have limited value for a North American user. But for persons working in this field, or for those planning to spend time in Europe, this will be a valuable addition to the already wonderful range of references available to them. Now if only someone would attack some of our continent’s problem species with equal thoroughness and vigour . . .! CLIVE E. GOODWIN 1 Queen Street, Suite 401, Cobourg, Ontario K9A 1M8 Canada north coast of Australia. This represents over one- third of the 242 mammal species known from the country, making it a very important group from the standpoint of mammal biodiversity. The field guide begins with a general introduction to the bats of Papua New Guinea followed by a chapter outlining the format of the book. There is a glossary for bio- logical terms encountered in the text, and an identifi- cation key to species. The bulk of the guide is devot- ed to the detailed species accounts for all of the bats found in the country. The accounts are separated into two chapters representing the two suborders of bats (Chiroptera). Within the Megachiroptera suborder there is only one family of Old World fruit bats or flying foxes (Pteropodidae), which is further subdi- vided into three subfamilies. The last chapter is on the Microchiroptera and it is split into eight sections representing the different families, or subfamilies for vespertilionid bats, found in Papua New Guinea. There is an useful appendix of latitudes and longi- tudes for the 713 localities mentioned in the book, which is followed by a very thorough bibliography for bats found in this region. The field guide ends with 24 beautifully illustrated colour plates of 58 bat species, 16 head and 42 full-body views, by Fiona Reid, and a checklist of the 91 species to keep track of your own personal life list. 338 Each species account begins at the top of a new page, which adds to the thickness of the book but speeds-up the locating of accounts, with the scientif- ic and common name. If there is a colour illustration, the plate and page number is referenced. Detailed information is provided in four descriptive sections on species identification, geographic range, natural history, and conservation status. One inconvenient drawback is that the distribution maps are grouped at the beginning of each family or subfamily without any page reference in the species account, or vice versa. The last three sections list the localities where the bats have been found, specimens examined by the author, and typical external measurements taken by field biologists. Where available, measurements are separated by sex and age, which is useful infor- mation for understanding variation, but unfortunately the average value is not presented as part of the range of measurements. A Field Guide to the Birds of Peru By J. Clements and N. Shany. 2001. The Ibis Publishing Company, Temecula, California. xvii + 283 pp., illus. U.S.$60. Following closely on the heels of Birds of Ecuador Volume I (Ridgely and Greenfield 2001) I automati- cally viewed this book in comparison to this other guide. The immediate reaction is that Birds of Peru is a true field guide, not like the oversized Ecuador ver- sion. .Drue,;at23,6em,.%. la,em << 3.6m (9 xX 6” X 1.25”] it will not fit easily in a typical pocket but, it will go in your pack. Even more remarkably, the authors have crammed all the known birds of Peru into this one manageable volume. Such a huge number of species has never before been included in a single book. We also have to remember that skulk- ing look-alikes or difficult-to-see high canopy dwellers form a large proportion of the birds covered. To put this in a Canadian perspective it is like observ- ing a few hundred species of Yellow Rail type birds or sorting out finches at the top of 200-ft high trees (the average Canadian White Spruce is 80 ft). The authors have tried to include all the birds of Peru, including the most recent additions and hypo- thetical species. Indeed, one species of hummingbird may well be new to science (or a hybrid?). The authors have included species that have recently been split, such as Caqueta Seedeater (formerly lumped with Variable Seedeater) and Nazca Booby (formerly with Masked Booby) or lumped like the Variable Hawk. Yet they have retained the old Cuvier’s Toucan Ramphastos cuvieri instead of White-throated Toucan Ramphastos tucanus. To keep the book size under control the text is very brief. Generally only a single sentence giving THE CANADIAN FIELD-NATURALIST. Vol. 116 One noticeable error in publication is the appear- ance of a chart for frequency of bats at different ele- vations (Figure 1.6) instead of a chart for different weights. Notwithstanding the minor aforementioned criticisms, I would highly recommended this field guide to any tourist or biologist heading to Papua New Guinea with an interest in bats. This is one group of animals that has an aura of mystery and intrigue, but with this informative and. well-illustrat- ed field guide in hand, it will open up a whole new world of exploration to which most people would never be exposed. I look forward to more books in this series on tropical nature. BURTON K. LIM Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, MSS 2C6 Canada key features is followed by a line on distribution in Peru. World distribution is covered in a single phrase. Where there are difficult identification prob- lems there is additional text; with some of the eupho- nias and terns for example. There are three maps; one on life zones (in colour), one on protected areas, and a general map. The first two are useful, but you would need a better road map to find the places named in the text. The authors often name specific localities for special birds, but the reader is on his own to find the exact whereabouts (e.g., the Saffron Siskin is at Tumbes). The authors also use the terms parano, puna, varzea, and terra firme throughout. I could not find an explanation of the meaning .of these important ecological divisions. This guide is, like the Ecuadorian one, another important step in bird literature from a neglected cor- ner of the world. Peru is home to 1800 species of birds, an incredible diversity. So this guide is most welcome. The authors were faced with covering a huge number of species while keeping the book to a manageable size. In this they have been most suc- cessful. It did, however, mean a number of compro- mises. The text gives only the key pieces of informa- tion on identifying marks, habitat and range. The only biometric given for virtually all the birds is length. There is no attempt to cover song but the cat- alogue number of an appropriate recording from the American Birding Association’s sales catalogue is ‘given (this means you would have to buy 30 CDs or tapes to cover all the birds). My copy has a photocopy insert listing the over 80 errors in the book. None of these errors are major but they are irritating. For example, the text omits all the references for the plate showing birds of prey in 2002 flight (16 errors) and for 27 species the plate does not give the text reference. The authors state that this was due to problems during printing on 11 September. However, these are not errors of printing but of proof reading and editing. Personally I would have preferred to delay the printing, rather than mar the impact of the new book. I also found a few addi- tional very minor errors. The illustrations are provided by a group of artists, with individual variation in style. For a large propor- tion the simplified style is reminiscent of the original Peterson, a style that works well for a guide. While the overall quality is good there are some problem areas. Some of the birds (Black-browed Albatross, and the boobies for example,) are oddly proportioned suggesting a lack of familiarity of the bird in the field. The colour of some birds (American Oystercatcher, Willet, Cock-of-the-rock, Pied Lapwing) is anomalous. Birds of Ecuador has 16 to 17 species illustrated on each plate, compared to about 14 in Birds of Peru. Yet the individual illustra- tions in Birds of Ecuador are much larger due to the more imaginative use of space. Birds of Peru plates have a large amount of white space. This makes a major difference in the detail shown. For example, The Blue-and yellow Macaw is almost 3'/2 times larger in Birds of Ecuador. Another factor I found confusing is that the artists have not been careful about scale. The Blue-and-yellow Macaw is depicted as smaller than the Red-bellied Macaw when it is actually well over 30% larger. I noted this on other pages too; the Spotted Sandpiper is about 10% undersized compared to the nearby Ruddy Turnstone and the Hoatzin is both small in depiction and under- sized relative to a neighbouring Cattle Egret. The plates do not always follow in sequence. The most widely separated species are the terns, being on plates 9 and 23. The black phase of the Vermilion Flycatcher has been put on a separate plate from the Birds of Delaware By G.K. Hess, R.L. West, M.V. Barnhill, and L.M. Fleming. 2000. University of Pittsburgh Press, Pittsburgh. 635 pp., illus. U.S.$65. When I opened the box in which this book was shipped, I immediately thought, “Wow!” This book, judged by its cover (and heft) is wonderful, and will make bibliophiles ooze with delight. This is not just an atlas of breeding birds, this is much more. This book incorporates data from many surveys, including the Breeding Bird Survey, Breeding Bird Census, Spring Migration Counts, Christmas Bird Counts, and of course, atlasing work from 1983 — 1987. In addition, data from museum BOOK REVIEWS 339 common red form. I could have understood this if it were on the page with the other small black flycatch- ers, but it is with several species of yellow-bellied flycatchers. Overall, however, the illustrations will work well for the intended purpose of this book, and that is to be a field guide In a book where space was obviously an issue, I am surprised they used so little of the pages facing the plates. The authors use between 25% and 50% of the space to give the English and scientific names plus the page reference. They could have put the sin- gle line giving distribution information on this page instead of in the main text, thereby cutting (or free- ing) 40 to 50 pages. Incidentally, repeating the scien- tific names in the text added about 10 pages, enough for a Spanish index. For, although the authors give all the names in Spanish, they do not include a Spanish index. It seems that there is more variation in the Spanish common names than there is in the English. Take stilt for example. Depending on the authority you follow the Common or Black-necked (Himantopus himantopus) or Black-winged Sult (4. mexicanus or H. h. mexicanus) has at least five com- mon names in the Spanish speaking world. The authors give those most widely used in Peru. None of my comments seriously detract from the major contribution the authors have made to ornithology. This is a valuable, useable field guide that will work well in the field and is a must-buy for Peruvians, visitors to Peru, and indeed birders in adjacent countries. Literature Cited Ridgely, Robert and Paul Greenfield. 2001. The Birds of Ecuador — Status Distribution and Taxonomy (Volume I), Field Guide. Cornell University Press. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Gloucester, Ontario K1J 6K5 specimens, government and non-government agen- cies, banding data, nest records are all incorporated. Field records from many individuals, some dating back many years, are also included. In short, with all these sources tapped, this book is a wealth of infor- mation that seems to cover all of the aspects of bird distribution and abundance that is available. Five introductory chapters precede the species accounts. The typical geology/climate and methodol- ogy chapters are there; a chapter describing historic records (from the 1630s!), which then follows the progression ornithology in the state, was an interest- ing read. Although there are plenty of maps in these — me ee eo eee re oa ee pe EE = — 340 chapters, including one map for each of the three counties' highways and population centers, I still felt that one map was missing. A single-page map show- ing key urban areas, important driving routes, and the frequently birded areas would have been extremely helpful to people not from the state. Every time I read of a birding site, I had to search the three county maps for the names for which I was looking. Twelve essays, dispersed at key locations through- out the book, bring an interesting perspective to Delaware's birds. There are essays on the history of Breeding Bird Atlases, the largest heronry in the state (Pea Patch Island), the spring shorebird migra- tion, laws concerning birds, and eight others. I liked this idea a lot. Line drawings (except for the most uncommon species) start each species account. These were drawn by a variety of artists, of a variety of skill lev- els. I can't understand why a few species are illus- trated only by a fledgling, not by an adult. The text briefly describes each bird's habitat, but most of the THE CANADIAN FIELD-NATURALIST. Vol. 116 text is devoted to analyses of migration dates, breed- ing information, and its distributional history in Delaware. A good amount of text is devoted to the atlasing data, and trends from the various surveys that occur throughout the year. For breeding species, a map is shown with squares in which breeding evi- dence was found; a second map is often included which shows the relative abundance of the species. CBC data (for the state's five long-running circles) are thoroughly summarized in a table and two graphs; these tables and graphs show absolute num- bers found and the more meaningful birds per party hour. All in all, this is a thorough work. It should be a model upon which other "Birds of ..." books should be based. RANDY F. LAUFF Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5 Canada Checklist of the Birds of Northern South America By Clemencia Rodner, Miguel Lentino and Robin Restall. 2000. Yale University Press, New Haven, CT. 136 pp., U.S.$23.50. This is a listing of 2245 species recorded in the strip of countries that run from Ecuador to French Guyana across the northern edge of South America. The coverage includes the offshore islands of Aruba, Curacao, Bonaire, Trinidad and Tobago. It does not include the Galapagos, Isla de San Andres, or Isla de Aves. The book is set out in tabular form. The listing gives each bird species by its most commonly used English name and its scientific name. There are no Spanish names given. The total number of sub- species worldwide is followed a summary of those subspecies found in the area covered by the book. There is an indication of the general locality of each subspecies The authors have developed a code for the altitudinal range and have a column for general comments (largely for site vagrant records and criti- cal species status). The final set of columns denotes the presence or absence of each species in the indi- vidual countries covered. The list is intended to be a companion to A Field Guide to the Birds of Northern South America, not yet published. The author’s intent is to provide a definitive list of species and names. They have also noted those subspecies that may be split in the near future. I was pleased to note that a number of the more recent changes in nomenclature have been included. The Common Piping Guan is split from Blue-throat- ed and Trinidad, the Golden —plumed Parrot is cor- rectly placed in the genus Leptosittica, Pterglossus erythropygius and P. sanguinneus are separate, and so on. However, I was surprised to find Buteo polysoma and B. peocilochrous had not been lumped (this took place a few years ago). The genus name Stercorarius has not replaced Catheracta, and Trogon viridus had not been separated from T. chionurius. The authors did not mention the potential split of Podiceps occip- italus juninnensis as the Northern Silvery Grebe. Glaucidium peruanum is still listed as the Peru Pygmy Owl not the more accurate Pacific Pygmy Owl. These are only a few examples of areas where I was disappointed that the authors had chosen to ignore changes that had been well accepted. I recog- nize this is a topic fraught with differences of opin- ion and revision, however I felt they could have been more consistent with emerging practice. I will await the publication of the field guide before deciding on the usefulness of this publication. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Gloucester, Ontario K1J 6K5 Canada 2002 BOOK REVIEWS 341 Birds of Southern South America and Antarctica By M. de la Pena and M. Rumboll. 2000. Princeton University Press, Princeton. 304 pp., illus. U.S.$24.95. This was originally published in England in 1998 by Harper Collins as Collins Illustrated Checklist - Birds of Southern South America and Antarctica. This guide covers over 1000 species in Southern Bolivia, Southern Brazil, Paraguay, Chile, Argentina, and northern Antarctica. (central Antarctica is not covered). All birds are illustrated in colour and accompanied by a cryptic text. Range maps are pro- vided at the end of the book. The bird’s name is given in English and Spanish or Portuguese as appro- priate. However not all the names are up-to-date. For example the “Imperial” Shag complex is out of date (also they are incorrectly called cormorants instead of shags). The Skua complex is in worse shape. The range maps also need a review. The Royal Albatross range is optimistic while that of the Emperor Penguin is extremely optimistic. The range for Turkey Vultures does not include the Falkland Islands, where they are common. I purchased this book as soon as it came out and used it in both South America and Antarctica. Although it looked like a perfectly good field guide I was disappointed. Several times I was unable to iden- tify a bird species using this book. The illustrations and information were simply inadequate to be cer- tain. On some, but not all, occasions I was able to dis- cuss my problem with an expert or use other texts to resolve the identification. However, I still have two, fairly distinctive birds that I have yet to identify. I was hoping that this reprint would also be a new version. This is not the case. All that has changed is the cover illustrations and the title. This is an oppor- tunity missed because the original idea was excellent and a thorough upgrade would have improved its usefulness immensely. The handy size and the great coverage would make it a very useful field book if it were a little more accurate. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Gloucester, Ontario K1J 6K5 Canada Whales and Other Marine Mammals of British Columbia and Alaska By T. Eder and I. Sheldon (with technical contributions by D. Pattie). Lone Pine Publishing, Edmonton. 160 pp. $16.95. How many sea mammal field guides do we need for the Northwest Pacific? This field guide alone quotes already four other field guides, and many more can be found on the market. However, this book by Tamara Eder and Ian Sheldon truly justifies its publication. It is nicely written and covers a huge amount of information within the full scope of a highly efficient field guide. Twenty-six whale species occur off British Columbia and Alaska, but the most prominent species are Grey Whale, Humpback, Orca, Minke Whale, and Harbour Porpoise. For an easy identifi- cation, the distinct diving sequences of eleven whale Species are provided. For many species, the book shows also major behaviour types (with drawings) to be observed by the whale watcher. The authors provide well structured sea mammal species accounts, even providing some subspecies information, which is normally not found in other classical field guides. The book considers not only British Columbia but all of Alaska. The superb draw- ings and photos (for seals) will be appreciated by the reader. But still, even with all this information provid- ed I find Saddleback Dolphins still very hard to distin- guish from the Striped Dolphin. Certainly, the sub- species of Saddleback Dolphins, (e.g., Short-beaked and Long-beaked) are still not distinguishable with this field guide, nor with any other. The following species are also very difficult to distinguish: Short- finned Pilot Whale, Risso’s Dolphin, False Killer Whale, Northern Right Whale Dolphin, Beaked Whales (Baird’s, Cuviers’, Hubbs’s, Stejneger’s), and Dwarf Sperm Whale. Perhaps new field guides and revisions should address these issues specifically in order to advance the knowledge for these species and their conservation? After reading this field guide the reader will be perfect in all the aspects of “Whale Watcher Talk”: The ancient Greeks had already a detailed knowl- edge and close relation with sea mammals; 70% of the global surface is covered by water; humans are not alone in the world; Endangerment Status of sea mammals according to USFWS and COSEWIC; Ambulocetus and Pakicetus whale fossils; character- istics and function of baleen; bubble-netting; lunge- and skim-feeding; biomagnetism; sounds and songs; dialects; diving skills; echolocation; whale strand- ings; intelligence of whales; relation between whales and natives; public perception of whales (from Moby Dick to Star Wars); dead Belugas treated as toxic waste; and the positive conservation effects of the U.S. Marine Mammals Protection Act. Besides clas- sical “Whale Watcher Talk” and whale records the reader learns in this book also about the value of whale fossils, that ecologically precious Ringed Seal fossils have been found near Ottawa, and that Bowhead whales can live over 200 years. 342 This field guide raises some interesting points: Are whales for entertainment or for profit? Whale watching along the West Coast creates approximate- ly 5 million $/year, plus 2 million $ for accessory items. With such intense, if not exploitive, whale watching activities, rules are important. As the authors claim, Alaska has the best and most stringent rules to make whale watching safe for the animals. In terms of “true” exploitation, the IWC (Inter- national Whaling Commission) and the whaling topic is also covered by the authors in detail; e.g., world-wide, as well as for the West Coast. Besides sea mammal related information, the authors cover environmental concerns as well. The ocean gets used as a dumping ground, and besides many others, “plastic bag” pollution contributes greatly to sea mammal deaths by blocking their digestive system; the authors quote “...that millions, or may be billions of plastic bags are discarded every day, many of which will end up floating in the Guia de las Aves de Espana By Eduardo de Juana y Juana Varela. 2000. Lynx Edicions, Barcelona, Spain, 223 pp., illus. 15 euros. This book is written entirely in Spanish, except that all common bird names are also given in English. It covers all the normally occurring bird species in Spain, and its offshore islands. It is laid out in modern field guide fashion with text on the left and illustra- tions on the right. Accidental visitors are placed in a section at the back of the book. Please note the English names are European not North American; e.g., Diver not Loon. Frequently the qualifier we nor- mally use is dropped; e.g., Black-crowned Night Heron is simply Night Heron. Oddly the Spotless Starling is listed as “Starling” while the common Starling of Europe and America is also given the same name. Surprisingly Sabine’s Gull is given as Larus sabini instead of Xema sabini. In addition, I noted that where the same or a similar species occurs in South America such as Great Egret, Skua, and Stilt there was hardly any conformance in the Spanish names. The text is well organized and provides the typi- cal information needed in the field. Included within the text are range maps giving summer, winter, and migration locations. The range maps include the immediately adjacent areas in southern France and northern Africa. A general map identifies the loca- tion of the top 50 areas to visit. Those people who do not know any Spanish at all will find the maps and illustrations useful. If you are willing to learn a few words (the colours, the seasons, and bird body parts primarily) the text is logical enough to follow the sense in general. Even a limited knowledge of Spanish will suffice to allow you to get a lot out of THE CANADIAN FIELD-NATURALIST. Vol. 116 oceans”! In addition, the authors state that the Northern Sea Lions are declining, and so are Harbour Porpoises. As the author suggests “Rare sightings are worth reporting” but unfortunately, the book does not tell us really how: no survey protocols or suggestions what to report are given. Besides the great text and design, one major drawback comes with the other- wise splendid book: it is poorly bound (glue); in my copy some pages fell out quickly after two days of usage. Here the publishers really need to re-think their decisions. Nevertheless, I find the authors have produced a very informative and useful field guide, which is lightly and well-written in a very appealing way. FALK HUETTMANN Geography Department.(Earth Science), 2500 University Drive N.W., University of Calgary, Calgary Alberta T2N 1N4 the text. The songs (“Canto” in Spanish) are ono- matopoeic, so are easy to follow. The authors include the current best estimate of population. This is a great innovation that I hope others will copy. It allows you to understand the differences in viewability with, say Bearded, Egyptian and Griffon vultures. The illustrations are well done and accurate. These include a rendition of juvenile, summer or winter plumages and subspecies where these are meaning- ful. Some juveniles are not depicted, notably some of the gulls. I was particularly impressed by the shape and attitude portrayed. It showed the artists were familiar with the bird in the field. The tougher groups (the warblers, shorebirds, pipits, and larks) are perhaps the best test and I had little trouble rec- ognizing almost all species. I only had trouble with a couple of localized species with which I am not familiar. The 113 accidental species that have occurred in Spain less than 20 or so times are recorded at the back of the book, but only 36 are illustrated. (The Glaucous Gull has been seen 63 times, but most of the other species are limited to one or two sightings.) I was surprised to see that Common Redpoll (Pardillo sizerin) has only been found once, in 1993. This is a wonderful, little, pocket-sized book that is well written and accurately illustrated. It will be most useful to residents of Spain and nearby commu- nities and to all visitors. Este libro esta escrito enteramente en espanol con excepcion de los nombres comunes de pajaros que también estah en inglés. Cubre todas las especies de 2002 pajaros que occuren normalmente en Espafia y sus islas costeras. Esta presentado como una moderna guia de campo, con el texto a la izquierda y las ilustraciones a la derecha. Especies accidentales aparecen en la parte posterior del libro. Este es un excellente libro, tamano bosillo, muy bien redactado y con ilustraciones veriidicas. Sera muy Util para los BOTANY Magical Mushrooms, Mischievous Molds By George W. Hudler. 1998. Princeton University Press, Princeton, New Jersey. 248 pp., illus. U.S.$14.95. This is an easy to read, nontechnical book that focuses on the effects fungi have and are having on man. The names Mushrooms and Molds refer to two specific groups of fungi. The facts are often present- ed with a good deal of humor and there is a mini- mum of technical terms. There are chapters on what makes up a fungus and how they are named; what fungi do and how they do it, that is how they grow, how they get nourishment, what enzymes they pro- duce to break down their food, how they fruit (pro- duce the mushrooms), and how spores develop. What fungi use as food is a principal theme because of the losses to humans when fungi consume (eat) plants, grains, etc. that we grow for food. Several chapters deal with fungi that damage or destroy our food and the forests. Brief but interesting discourses treat the Potato Famine in Ireland and its effects on North American demographics, the Powdery Mildew of Grapes that nearly destroyed the wine industry in Europe, and the Rust diseases of wheat and their potential impact on the world’s food supply. Also discussed are catastrophic tree diseases, such as Chestnut Blight, Dutch Elm Disease, and White Pine Blister Rust, that resulted when spares of fungi were inadvertently brought into North America by the activities of man. An entire chapter is devoted to Ergot, a disease of cereals and grasses, not because it is a major problem but to explain the vari- ous ways the fungus has influenced man. Eating the bread from such flour contaminated with Ergot can cause poisoning ranging in seriousness from halluci- nations to loss of limbs and agonizing death. The cause of the Salem Witch Trials and the gruesome BOOK REVIEWS 343 residentes en Espafia y las comunidades vecinas, y también para los turistas. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Gloucester, Ontario K1J 6K5 Canada disease of man called St Anthony’s Fire have been attributed to Ergot poisonings. Additional topics that are the subject of individual chapters are mycotox- ins, mycoses, medicinal molds, yeasts for baking and brewing, edible and poisonous mushrooms, hallu- cinogenic mushrooms, wood decay, fungi and insects, and symbiotic relationships between fungi and plants. Finally, a few critical comments. On page 200 it says the Dry Rot Fungus (Serpuls lacrimans) does not occur in the western hemisphere, but a 1993 reference lists it from 37 States and 7 Provinces. Although dry rot is widespread in North America, poured concrete foundations that are waterproofed, and central heating prevent Dry Rot Fungus from becoming a major prob- lem. Although the color pictures are very good, the quality of the black-and-white figures varies greatly. The drawing (Figure 10.3) labeled Hericium eri- naceus is not that fungus but H. ramsoum. And the title of the associated paragraph (page 162) would be more accurate if the word “erinaceus” is changed to “species” because there are four Hericium species in North America, all are edible and several are more commonly found that H. erinaceus. Most naturalists recognize a few mushrooms but know little more of the fungi. Professor Hudler’s book provides a broad view of the fungi and is a good beginning toward understanding how fungi impact birds, wildflowers, trees, and many other aspects of the natural world. Jim GINNS 1970 Sutherland Road, Penticton, British Columbia V2A 8TS8 Canada 344 Spatial Pattern Analysis in Plant Ecology By M.R. T. Dale. 1999. Cambridge University Press, New York. 326 pp., illus. Cloth U.S.$69.95; paper U.S.$35.95. As the title of this book aptly suggests, Spatial Pattern Analysis in Plant Ecology is a review of techniques used by ecologists to study spatial pat- terns of plant populations. The book has nine chap- ters, covering a wide spectrum of techniques. The first two chapters are introductory, outlining con- cepts that underlie spatial analysis and sampling techniques. Titles of the remaining chapters illumi- nate their contents: Basic methods for one dimension and one species, Spatial pattern of two species, Multispecies pattern, Two-dimensional analysis of spatial pattern, Point patterns, Pattern on an environ- mental gradient, and finally, a concluding chapter. Techniques described throughout these chapters include semivariograms, correlograms, fractal analy- sis, spectral analysis, and blocked, paired, and local quadrat variance analysis, among others. In the preface, Dale states that the book is designed to help the reader understand the concepts and meth- ods of spatial pattern analysis. Spatial Pattern Analysis in Plant Ecology is part of the Cambridge Studies in Ecology series, which presents up-to-date reviews of topics within ecology, and is aimed at “advanced final- year undergraduates, graduate students, researchers, and university teachers, as well as ecologists in indus- try and government research”. Dale’s book certainly meets this standard, although I would add a caveat that the greatest benefit will be obtained by readers who have an understanding of statistical methods but are reasonably new to the world of spatial analysis tech- niques. Dale presents a good, thorough review of tech- Rare Vascular Plants of Alberta By Linda Kershaw, Joyce Gould, Derek Johnson, and Jane Lancaster. 2001. The University of Alberta Press and The Canadian Forest Service, Edmonton, Alberta xliv +484 pp., illus. $29.95. This is a fascinating book which will catch the eyes of many individuals who have previously not paid much attention to the plants that grow around them. In it is information about 485 species which are of rare occurrence in the province of Alberta, a large percent- age of the total known to occur there. The book is divided into sections; an introduction which is fol- lowed by the treatments of Trees and Shrubs, Monocots, Dicots, Grass-like Plants, Ferns and Fern Allies, Addendum (which includes a few plants found since the main text was completed in 1998 and two deletions), Appendix | (Keys to rare Botrychium and Isoetes found in Alberta), Appendix 2 (Rare vascular plants of Alberta by Natural Region), Appendix 3 THE CANADIAN FIELD-NATURALIST Vol. 116 niques. However, rather than an in-depth description of particular techniques, the book contains an intro- duction to a broad range of material. In each chapter, the basic logic and fundamentals of a technique are described, often, but not always, including equations. This description is usually supported with examples from the literature, and citations of sources to which the reader could refer for more details. The figures are simple and supportive of the concepts in the text. There are few worked examples. Although Spatial Pattern Analysis in Plant Ecol- ogy is aimed at plant ecologists, the methods described are applicable to a range of systems and taxa. Thus, its scope is broader than plant ecology. The more traditional analysis methods, such as blocked, paired, and local quadrat variance analyses, are covered in greater detail within this book, than are more novel techniques, such as geostatistics, wavelet analysis, landscape metrics, and fractal anal- ysis. Indeed, promising techniques such as lacunarity analysis and variance partitioning ordinations receive only passing references, and tree regressions are not mentioned at all. On balance though, the book is full of good ideas and methods and, because of the referenced examples, is a fine starting point for those who are new to spatial analysis. I recom- mend Spatial Pattern Analysis in Plant Ecology for such an audience. JEFF BOWMAN Wildlife Research and Development Section, Ontario Ministry of Natural Resources, 300 Water Street, 3rd Floor North, Peterborough, Ontario. K9J 8M5 Canada; e-mail: jeff.bowman @mnr.gov.on.ca (Species in this guide that are not found in the Flora of Alberta (second edition) — New Discoveries and changes resulting from taxonomic revision), Appendix 4 (Taxa previously reported as rare for Alberta but not included in this book), Appendix 5 (Species reported from or expected to occur in Alberta but not yet verified), Appendix 6 (Alberta taxa classi- fied as nationally rare by Argus and Pryer (1990)), Appendix 7 (Rare native plant report form), Glossary, References, Index and Photo Credits. In the main text most pages include information on only one rare plant: common name, scientific .name, family name, descriptive information, habi- tat, and notes of special interest. On each page the colour photographs, line drawings, coloured distri- bution maps, and maps of North America which depict other provinces where a species is rare or common are all most important and make this book 2002 BOOK REVIEWS 345 so very attractive. The four editors and their many assistants are to be congratulated. WILLIAM J. CoDYy ENVIRONMENT Biological Resources Program, Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada. The Global Environment in the Twenty-First Century: Prospects for International Cooperation Edited by Pamela S. Chasek. 2000. United Nations University Press, Tokyo. xi+465 pp., illus. U.S.$39.95. This book focusses not on the biological environ- ment which naturalists love but on the political and economic challenges facing its survival. The contribu- tors are international experts in environmental policy and its bureaucratic aspects. The piecemeal progress on environmental issues is examined through the roles of five types of actors. Under states and their sovereignty, the review includes energy in India and China (especially, of course, the Three Gorges Dam), issues before third-world cities (with Brazilian exam- ples of successes, such as as Curitiba’s infrastructure, and failures, such as Rio’s pollution agency), and tem- poral trends such as leapfrogging technological inno- vations to new forms like photovoltaics. Under civil society, and its reflection in non-governmental organi- zations, there is much fascinating material. The power, presence, and alliances of these sovereignty-free “views from nowhere” are demonstrated in a range of impacts including waste treatment by states, disposal of industrial oil rigs, and social mores such as the Canadian hunt on harp seals. In terms of accountability and performance in relation to governments and indus- try, of advocacy and service, and of drawing on sci- ence, non-governmental organisations (NGOs) are the stars of the show. By contrast, under markets, the dis- astrous consequences of unconstrained activity dictate the need for meta-market vision and knowledge, glo- bal governance, and full-cost pricing. Canadians will find the chapters on freshwater, in which it is argued that it must be given a price, and on agriculture, much changed under the new GATT, most interesting. Under regional arrangements, efforts to cope with massive problems, especially in Africa, Asia, and Eastern Europe, are considered. Issues range from loss of natural habitat to the impact of “trade, not aid” poli- cies. Under international organizations the focus is on track records of the World Bank and the UN Environmental Program (UNEP) and Commission on Sustainable Development. In the Conclusion Chasek provides a synthesis of the dominant themes of public attention, governance, conflicts, and flawed markets, to extract some potentially fruitful directions. The chapters are well documented, with a happy balance of description and analysis. Figures and tables are used effectively, although some chapters give More Than You Ever Wanted To Know. The even- ness of discussion does credit to the editor. The four pages of collected acronyms are invaluable. Would it not have been indicative of the new awareness of environmental linkages to use economic “externality” in quotes? It is good to see dialogue between chapters, often lacking in multi-authored works. For instance, whether UNEP scaled up to some sort of World Environmental Organization would solve certain existing problems is argued between chapters. As a product of the UN, the book pays particular attention to the current and potential roles of that organization and its agencies, as well as to topics and areas beyond the first-world. And surely the world must move from soft motherhood declarations to hard agreements with teeth. This book provides an in-depth examination of the international structures and dynamics for such environmental policy. The contrast between actual crises, natural and human, of conser- vation and education, and the glossy bureaucracy intended to deal with them is stark. Against the good news, such as the acceptance that national sovereignty should not prevent joint action, is a much larger stack of bad on the Aral Sea, the African Matatuine District, and on and on. Most valuable in this book are the rec- ommendations, in various chapters and the Conclusion, which outline possible courses of action. The set of existing agencies and treaties is already labyrinthine, and it is difficult to see how to add effectively to it. Much more deeply worrying is the impression that the entire apparatus is functioning, and can only function, as reporters and cheerleaders while the environmental destruction continues apace. PATRICK COLGAN Royal Botanical Gardens, POB 399, Hamilton, Ontario L8N 3H8 Canada eee ee ee ee eS 346 Reading the Entrails: An Alberta Ecohistory By Norman C. Conrad. 1999. University of Alberta Press, Calgary. xviii + 197 pp. $19.95. Reading the Entrails is best described as a good rant. Beginning with the Wisconsinan glaciation and carrying the story through to the reign of the Albertan Conservatives and Ralph Klein, Norman Conrad pro- vides an intensely personal look at the ecological his- tory of the province and the irreversible damage that has been wreaked in only a few generations. His spir- ited — some might say, relentless — style makes for a provocative read. But it is also an uneven, at times frustrating, analysis. Conrad makes some questionable statements about the coming of agriculture to the western interi- or in the nineteenth century. He suggests, for exam- ple, that small-scale commercial farming was already underway in the early 1800s, that the fur trade ended in 1870 with the transfer of Rupert’s Land to Canada, and that First Nations did not understand the treaties and were “effectively elimi- nated” as a people in a few decades. His assessment Spatial Optimization for Managed Ecosystems By J. Hof and M. Bevers. 1999. Columbia University Press, New York, NY. 258 pp., illus. Cloth U.S.$65 in U.S.A., U.S.$75 elsewhere; paper U.S.$30 in U.S.A., U.S.$34.50 elsewhere Spatial Optimization for Managed Ecosystems 1s part of a series called Complexity in Ecological Systems. Other well-known titles from this series include Toward a Unified Ecology (Allen and Hoekstra 1992) and Ecological Scale (Peterson and Parker 1998). With Spatial Optimization, Hof and Bevers demonstrate the use of mathematically pro- grammed optimization models to determine strate- gies for optimal allocation of resource management interventions. Optimization models have much in common with simulation models, but optimizations are different in that they are intended to accommo- date and reduce a large number of possible scenarios to a set of manageable options. Whereas simulation models frequently are developed to deal with spatial structure, few optimizations methods accommodate spatial considerations. This book is a novel demon- stration of the potential utility of spatial optimiza- tion. The book is divided in four parts: (1) Static spa- tial relationships; (2) Spatial autocorrelation; (3) Dynamic movement; and (4) Diversity and sustain- ability. Within each of these sections, information is presented in a modular format, where similarly structured modules present spatial optimization approaches to a range of increasingly complicated management problems. Optimization models that THE CANADIAN FIELD-NATURALIST Vol. 116 of the region’s development during this period is also firmly fixed on the open grasslands; his gaze rarely extends north to the boreal forest and activities there in the nineteenth century. Conrad is much better in describing developments in the post-World War Two period, especially in the provincial north. He raises some fundamental ques- tions about the nature and impact of Alberta’s petro- chemical and forestry industries. He also wonders aloud whether society can really afford the costs of a resource-based economy in the long run. Clearly, the environment cannot; according to Conrad, at the cur- rent pace, there will be nothing left to do in the near future but “read the entrails” of a once richly diverse environment. And that’s why this book should be read by anyone seeking some alternative perspective on Alberta’s relationship with the land. BILL WAISER University of Saskatchewan, Saskatoon, Saskatchewan, S7N 5A5 Canada use integer and non-linear programming are used in all scenarios to find the optimal solution, where optimal is defined as an approach that meets all stated management constraints simultaneously. Management problems presented are diverse, and include such things as wildife-habitat relationships, timber harvesting, pest control, and water runoff. Regardless of this diversity however, the same modular approach is taken to each problem. This presentation successfully demonstrates optimiza- tion, as it becomes obvious to the reader that a simi- lar spatial optimization approach can be applied to a wide range of management problems. A number of interesting examples are provided, such as opti- mizing Black-footed Ferret, Mustela nigripes, rein- troduction in South Dakota. These provide useful context for the reader. Although the approaches outlined in Spatial Optimization have potential for great utility to resource managers, readers are faced with some challenges in processing the material. Foremost, as with any modelling exercise, optimization models are contingent on a number of assumptions. These vary with the application but are of the typical sort, such as specifying the shape of dispersal distribu- . tions. Hof and Bevers are explicit about these assumptions, which is a strength of the book, but the reader will nonetheless note that decisions made in light of assumptions could be incorrect. Further, as complicated as these models are, they still oversim- plify systems, frequently omitting complicating 2002 dynamics such as competition or predator-prey inter- actions. Finally, readers of this book (and potential users of spatial optimization) will be faced with somewhat dense discussions of integer and non-lin- ear computer programming. The methods described in Spatial Optimization are challenging, but they have potential for great utility in solving management conflicts. As such, I recom- The Amber Forest By George Poinar Jr., and Roberta Poinar. Princeton University Press, Princeton, 2001. 239 pp., illus. U.8:$29.95. Historically, “the ancient Greek poets’, says early 20" century gemmologist George Kunz, “the grains of amber were [considered] the tears annual shed over the death of the brother Phaethon by the Heliades after grief had metamorphosed them into poplars growing on the banks of Eridanus” (Kunz 1913 [1971], pages 55-56). We now consider amber not only as artifacts of past cultures, but evidence of life long before humanity. The most prolific investi- gator of the contents of amber is George Poinar (see Quest for Life in Amber and Life in Amber), and with Roberta Poinar, their most recent, colorful, tome is The Amber Forest. The subtitle of this recent work, “a reconstruction of a vanished world”, is a geographically restricted examination of amber found on the Caribbean island of Hispaniola. The amber, however, has a broader range, covering a time period from 15 to 40 million years ago. Collecting these samples are amber min- ers. And the mines “are little more than small, tortu- ous tunnels carved into the sides of the mountains or sometimes pits sunk deep into the ground” (page 6). The rewards are often sold to the highest bidder. By examining the collections, both in museums and in private hands, the Poinars illustrate unique snapshots of the past; features we more often than not could never see by other methods of fossilization. The Amber Forest, however, is not a general read. It is a systematic look at the flora and fauna found in the fossilized sap. From plants and seeds, to insects (of all shapes, sizes, and types), to frogs, lizards and BOOK REVIEWS 347 mend the book for readers who have a serious inter- est in resource management. JEFF BOWMAN Wildlife Research and Development Section, Ontario Ministry of Natural Resources, 300 Water Street, 3rd Floor North, Peterborough, Ontario, K9J 8M5 Canada; e-mail: jeff.bowman @mnr.gov.on.ca snakes; all have left traces or whole bodies in amber. Who would not be moved at a specimen of Sphaero- dactylus, a gecko lizard, in amber, skin and all, with a partially chewed leaf. “Could the notch on the leave have been made by an insect the gecko attempted to seize?” ponder the authors. Literally or metaphorically “frozen” bodies often lead us to these kinds of thoughts and questions. Despite the group-by-group framework of this volume, the 171 black-and-white, and 171 colour photographs are spectacular. Kudos to nature for providing us with these examples of past life, and to the photographer for capturing these images. It is no wonder that many of these objects are often consid- ered works of art. Supporting the text and pho- tographs, are black-and-white illustrations of the inhabitants of this ancient tropical forest, as they appeared in life, as they interacted with other occu- pants of the same ecosystem. Unfortunately, we are unable to fully realize the wealth of information available in samples of amber. Many of the finer specimens are sold privately and are not seen by scientifically trained eyes. What we do know, what The Amber Forest vividly illustrates, is that the possibilities of unveiling nature of the past are seemingly endless. Literature Cited Kunz, G. F., 1913 [1971]. The Curious Lore of Precious Stones. Dover Publications, Inc., New York. 406 pages. TIM TOKARYK Box 163, Eastend, Saskatchewan SON OTO Canada The Natural History of an Arctic Oil Field, Development and the Biota Edited by Joe C. Truett and Stephen R. Johnson. 2000. Academic Press, San Diego. 422 pp., illus. U.S.$ 69.95. “Oil Exploration and Development ... result in environmental change”. This excellent book describes and investigates these topics, but also fur- ther introduces and consolidates human activities and oil development in ecological research and natu- ral history; it sets new publication standards on this subject and polar research, e.g., to be followed in the Russian Arctic and elsewhere. For the Alaskan Arctic, industrial development and oil production started 50 years ago. In 1968, the 348 Atlantic Richfield Company announced the discov- ery of a major oil accumulation near Prudhoe Bay, Alaska. But already in 1969, the U.S. government passed the National Environmental Policy Act (NEPA) requiring the full disclosure of the environ- mental costs of major development ventures. This needs to be seen in the context of Prudhoe Bay being among the largest oil and gas discoveries in the his- tory of North America. Oil from the Alaskan Northern Slope contributes to the approximately 85% of the Alaskan state budget derived from taxes and royalties. Besides Prudhoe Bay having the oldest oil fields in Alaska’s Arctic, Prudhoe Bay also pro- vides over 20% of the USA oil production. Based on “Oil Field Research”, the book consists of 19 research papers (no abstracts) from 27 authors (contact addresses provided) which are mostly asso- ciated with universities or with international high- caliber research consultancies such as LGL Ltd. (including the editors) and others. Published by an international high-quality publisher (Academic Press), all of the papers are (externally) peer- reviewed except for the Introduction, North Slope Development and the Synthesis; this assures objec- tivity of the science presented. Research and publica- tion costs were carried in part by BP Exploration (Alaska) and others, who also set up the focus of the book and authors. This research publication describes almost all major aspects of wildlife (except for sea mammals, raptors and jaegers) related to oil development and production over the last 20 years and beyond. Divided into four parts, the chapters deal with fish, wildlife, communities, and habitats of the terrestrial and marine ecosystems. The book gives the reader a realistic overview about environmental assessment, industrial (Arctic) research, and governmental regu- lation and planning processes. It presents nicely and detailed the Environmental Training Programs required for (oil-) field personal, and it shows the mitigation measures which oil companies undertake. The “Boulder Patch’, also a location of major oil and gas reserves off Prudhoe Bay, is THE most diverse marine area in the Beaufort Sea. Due to the oil development the Prudhoe Bay region has finally received more research attention than any other and similar locations, e.g. Arctic Canada and certainly Newfoundland (Hibernia) and Gulf of Mexico. As with any oil impact study the monitoring, pre-spill, and pre-development data are very crucial, but rarely available; some are presented in this book. For instance, research in Prudhoe Bay has resulted in major biological, long-term data sets (hopefully made available soon to the global community via Internet/WWW): 18 years of fish data in the Beaufort Sea, 14 years of Snow Geese research, and what may be the only long-term data set on shore- bird migration. Prudhoe Bay has received a very THE CANADIAN FIELD-NATURALIST Vol. 116 detailed geo-botanical mapping, including the most extensive benthic sampling done anywhere in the Beaufort Sea. Nine years of long-term studies on fish populations in the study area allow for strong cap- ture-mark-recapture analysis and powerful results. Besides others, the study area has a wonderful long- term Polar Bear Satellite GPS study, and a very detailed description and summary of the Grizzly Bear population. Several Caribou herds exist in Alaska [e.g. Western Arctic Herd, Central Arctic Herd (CAH), Tesehkpuk Lake Herd, Porcupine Herd], but only the CAH is really located within the Prudhoe Bay Oil Field area. The Caribou papers pre- sent a great summary of Caribou research, and they are a solid resource for references. It is shown that Caribou numbers increased in recent years. They do use oilfields to avoid mosquito harassment; but the impact of roads and elevated pipelines on Caribou migration and biology is unclear and currently in debate. Changes brought by humans to the Arctic are nor- mally centred around (1) addition of gravel, (ii) dis- ruption of tundra surfaces, (111) creation of impound- ments, and (iv) introduction of elevated structures. Oil development clearly improves human access to wildlife, such as Grizzly Bears. If not regulated, this can result in increased (hunting) pressures and dis- turbances. The availability of garbage affects Arctic Foxes, which have high densities in the Prudhoe Bay area; a simple removal of these predators with traps remains doubtful. For the study area, the book reports dramatic increases of Black Brant in the 1980s, Red- necked Phalarope show high densities in impound- ments, and Grizzly Bears are more productive on oil fields (but the offspring is also more likely to be killed). Although the oil production area “P Pad”, built in the 1990s, is already much smaller than “A Pad” (built in the 1970s), much of the research pre- sented still deals with the gravel (road) effects. Impoundments have indeed more nutrients than natu- ral waterbodies; culverts can improve/restore fish migration. As a major impact of oil development, population increases are observed for several shore- bird species, Black Brant, Snow Goose, Tundra Swan, Caribou, Grizzly Bear, Polar Bear, and proba- bly Arctic Foxes; but Dunlins declined. However, the study area of Prudhoe Bay was defi- nitely not untouched and “virgin” before oil develop- ment occurred: Muskox were introduced, Polar Bears were hunted from the air between 1967 and 1974, strychnine was widely used to exterminate Wolves but also killed non-target animals such as . Grizzly Bears and Squirrels. An historic shorebird hunt existed all over North America, and Eider Ducks and other wildlife were hunted by natives. The papers of the book give excellent overviews or summaries, but sometimes they don’t show meth- ods in full detail. Some of the study objectives and 2002 BOOK REVIEWS 349 designs are somewhat worrisome for ecologists; for instance, studies presented have a strong focus on animal abundance and charismatic or huntable species. However, this appears to be in the interest of the general public. Low-level flying, viewshed effects, environmental ethics, and individual animal suffering due to oil development are not addressed. This might confirm that research designs were driv- en by questions directly of service to the oil industry, but not necessary designed to investigate purely eco- logical and natural history research questions. Some sample sizes interpreted are quite small, power-tests to evaluate population trend statements are missing, and DISTANCE sampling to survey abundances was never applied. Studies dealing with ecosystem ener- gy flux and predator-prey balances, (e.g., “explod- ing’ abundances of wildlife and predators), are somewhat lacking. Impoundment effects on ecosys- tems are unlikely to be fully understood with simple comparisons and experiments. Almost all papers include unpublished reports (grey literature) from over 20 years of consultant contracts, but perhaps they should be specifically quoted and marked in the research papers, as done elsewhere and in similar publications. The photos in this book are beautiful (some enlargements would have been excellent); for my taste, some maps are a little repetitive in their con- tent. Finally, it would have been nice to have an overview about research gaps, and urgently to be investigated conservation needs and threats for Prudhoe Bay, as well as a detailed list of all oil spills that have occurred. Although not all ecological questions are fully investigated in this book and are sometimes referred to as “unstudied”, some extremely important man- MISCELLANEOUS agement questions are raised in this publication land- mark: “The assumption of habitat limitation on the breeding grounds is not based on evidence, but rather is a conservative approach to shorebird con- servation given the lack of basic population informa- tion on most species involved”. That quote from the book is a classic statement valid for most migratory species, and might well summarize the current (lack of) knowledge in population studies. Here another “wildlife classic” raised in this book: does simple (bird) abundance equal (nesting) densities ? Obviously, and as the book suggests, consistent and meaningful survey methods for better science and management are required. Despite the major research efforts presented, the following citation might prove to show how weak our knowledge can be even after long-term studies: “We believe that the presence and activities of humans in the oil fields have had little or no population-level impact on species ..., although data currently are insufficient to confirm this belief”. Nevertheless, the authors can be congratulated for their outstanding product. A complete index, and an Errata with only six (!) minor corrections, comple- ment this excellent book and make it a perfect refer- ence, description and template for similar studies and publications. What will the future bring for Prudhoe Bay and its oil development endeavors? With such type of research effort and publication one could rest assured indeed. FALK HUETTMANN Centre for Wildlife Ecology, Biological Department, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia, V5A 1S6 Canada Finding Order in Nature: The Naturalist Tradition from Linnaeus to E. O. Wilson Paul L. Farber. 2000 Johns Hopkins Introductory Studies in the History of Science. Johns Hopkins University Press, Baltimore. 136 pp., illus. Cloth U.S.$39.95; paper U.S.$15.95. Paul Farber has done it again! In his latest book he tells succinctly how natural history has yielded the major unifying theory of the life sciences, uncovered some of the deepest insights into nature, led us to con- cern for the environment, and attracted public interest for more than two and a half centuries. Animals were named in Genesis but natural history did not emerge as a scientific subject until the eighteenth century. Farber begins with a Swede, Linnaeus, his con- ception of order, and his view of creation as a bal- anced and harmonious system. Linnaeus’s students, his “apostles,” amassed great collections. Although the Linnaean classification system was artificial, his binomial nomenclature became the universal world standard. A Frenchman, Buffon, shared the birth-year of 1707 with Linnaeus, but outlived him by ten years. He developed the Jardin du roi, in Paris, as the fore- most institution in its day for the study of the living world. Buffon’s 36-volume encyclopedia of animals was the second most frequently owned item in pri- vate libraries in France. Buffon gave less emphasis to classification than did Linnaeus. He considered nature, not God, to be the generative power. Collectors soon circled the globe to send back specimens to museums in major European cities. 350 Advances in paper manufacture and printing allowed artist-naturalists such as John Gould and John James Audubon to publish small editions of large, beautiful books. Scientific journals began and by about 1850 natural history had begun to divide into disciplines and sub-disciplines. A German, Alexander von Humboldt, emphasized measurement, visual representation, and the search for laws that dealt with nature. However, Paris remained the international hub of natural history through the first half of the nineteenth century. Saint-Hilaire studied comparative anatomy to demonstrate the similarities and differences between different animals. Cuvier described the mass migra- tions and mass extinctions that had occurred in the geological past. Charles Lucien Bonaparte ruled as the greatest expert in ornithology; he listed 7000 species worldwide. The last of the early pioneers was an Englishman, Charles Darwin. His grand unifying theory of evolu- tion brought together the disparate studies of classifi- cation, embryology, behavior, adaptation, morpholo- gy, paleontology, and distribution. The term biology, favoured by Huxley, gradually came into favour. Physiological, embryological and cell research, using the experimental method, became increasingly important. During “the Golden Age of Natural History, 1880, 1900,” zoos, museums and botanical gardens made natural history accessible to the general public. Visitors thronged to the new edifice housing the Excavation By Steve Roskams. 2001. Cambridge University Press, Cambridge, 311 pp. $43.50. The science of archaeology is a little over a hun- dred years old. Previous to the mid-19" century, archaeology, like many natural sciences, was often restricted to trophy hunting. Today, as a result of a century of investigation and development, archaeolo- gy seeks information of all kinds. The acquisition of said information requires a broad respect for avail- able resources and surrounding environments. Cambridge Manuals in. Archaeology has produced its 14" installment titled Excavation, and is the most recent summation of the acquisition process. University of York Lecturer Steve Roskams provides a well thought out series of steps involved with archaeological investigation, covering diverse possi- bilities, tools, and methodology. The fourteen chapters can be seen as covering three main sections, the first being preparation before entering the field, probably the most vital. Beginning with a short historical overview of the excavation process, Roskams highlights such basic THE CANADIAN FIELD-NATURALIST Vol. 116 British Museum of Natural History when it opened in South Kensington, London, in 1881. Similar “cathedrals of science” became available in most major cities. There were more than 200 botanical gardens, worldwide. The rediscovery of Mendel’s Laws in 1900, Dobzhansky’s use of chromosomes to divide Drosophila into two species, and Ernst Mayr’s bio- logical species concept led to studies of the genetics of population change. The final chapter tells of E. O. Wilson who stud- ied pheromones as signalling chemicals among insects. He became concerned with biodiversity, then wrote Sociobiology, and called for a return to the naturalist tradition, and a realization that natural his- tory is central to the life sciences. This recommenda- tion takes us back to where the book began, to Linnaeus and Buffon. Q.E.D. This inexpensive paperback is primarily directed to the student and to the general public. It provides insight into the background, motivation, and achievements of the key players, all male. It should be compulsory reading for any scholar entering a biology program. Well-read naturalists will find it a useful review; they will be pleasantly surprised by the many interesting facts unearthed by Farber in his well-written presentation. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N 0J8 Canada practices as test pits, remote sensing technology, to aerial photography. Reaching farther afield, the importance of consequential activities like finance and administration, safety in the field, to archival resources must be approached before any excavation can begin. The second section, chapters 5 to 9, encroaches on the process of excavation itself, centering on reasons and methods of record keeping including spatial and stratigraphic methodology. Drawings, photographs, grids, all help the field activity to “take place in con- trolled conditions, allowing full recording of the physical character and spatial disposition of the stratigraphic units in a site” (page 153). Without these measurable terms of reference, we slip back into trophy hunting. The final section, chapters 10 to 14, covers the act of collecting and interpretations based on physical evidence such as sedimentary deposits and how to approach non-deposit finds such as buildings, bod- ies, and artifacts. Accomplishing any recovery or discovery, “the stuff of archaeological research” 2002 BOOK REVIEWS 351 (page 266), in a consistent manner, helps clarify the evidence with regard to broader issues and theories within archaeology. Other factors, however, must also be considered such as politics, environment, ideology, and funding, which together, fall outside of the scope of the field worker yet can hardly be dis- missed in deciding where to excavate and what type of information is anticipated to be gained. No apologies are required for the primarily British-based examples provided in Excavation. NEw TITLES Zoology 7All-weather hawk watcher’s field journal. 1999. By D. S. Heintzelman. J. L. Darling Corporation, Tacoma, Washington 50 pp., U.S. $14.95. Awe for the tiger, love for the lamb: a chronicle of sen- sibility to animals. 2002. By R. Preece. UBC Press, Vancouver. 488 pp. $85. Birding across North America: a naturalist’s observa- tions. 2002. By P. E. Keenan. Timber Press, Oregon. 260 pp., illus. U.S. $29.95. *Conserving bird biodiversity: general principles and their application. 2002. Edited by K. Norris and D. J. Pain. Cambridge University Press, New York. xiii +337 pp., illus. Cloth U.S. $100; paper U.S. $38. +Firefly encyclopedia of insects and spiders. 2002. Edited by C. O’Toole. Firefly Books, Willowdale, Ontario. 240 pp., illus. $40. *Firefly encyclopedia of reptiles and amphibians. 2002. Edited by T. Halliday and K. Adler. Firefly Books, Willowdale, Ontario. 240 pp., illus. $40. *Fishes of Alaska. 2002. By C. W. Mecklenburg, T. A. Mecklenburg, and L. K. Thorsteinson. American Fisheries Society, Bethesda, Maryland. xxxvii + 1037pp + plates, illus. U.S. $129. *Fishes of the Gulf of Maine. 2002. Edited by B. B. Collette and G. Klein-MacPhee. 3" edition. Smithsonian Institute Press, Washington. xxxiv + 748pp., illus. U. S. $75. Game in the garden: a human history of wildlife in western Canada to 1940. 2002. By G. W. Colpitts. UBC Press, Vancouver. 216 pp., illus. $75. *Global register of migratory species (GROMS): data- base, GIS Maps, and threat analysis. 2002. By K. Riede. German Federal Agency for Nature Conservation. 404 pp. *Grassland grouse and their conservation. 2002. By P. A. Johnsgard. Smithsonian Institute Press, Washington Those working in other geographic regions (and those working in related fields like palaeontology) can still utilize the significant resources and method- ologies provided in this volume. Either as an “ama- teur” or “professional”, Excavation is a worthy companion for any excavator. TIM TOKARYK Box 163, Eastend, Saskatchewan, SON OTO (Canadian distributor Scholarly Book Services, Toronto). xiv + 151 pp., illus. $57.75. *Paleoimagery: the evolution of dinosaurs in art 2002. By. A. A. Debus and D. E. Debus. McFarland Publishers, Jefferson, North Carolina. 285 pp., illus. U.S. $49.95. A Passion for wildlife: the history of the Canadian Wildlife Service. 2002. By J. A. Burnett. UBC Press, Vancouver. 320 pp., illus. $85. +Sealworms in the north Atlantic: ecology and popula- tion dynamics. 2001. Edited by G. Desportes and G. McClelland. North Atlantic Marine Mammal Commis- sion, Tromso, Norway. 171 pp., illus. *Sibley’s birding basics. 2002. By D. A. Sibley. Knopf, New York. 154 pp., illus. U.S. $15.95; $23.95 in Canada. Wildlife viewing in North America: a management planning handbook. 2002. Edited by M. J. Manfredo. Oregon State University Press, Corvallis. 336 pp., illus. U.S. $24.95. +The lost dinosaurs of Egypt. 2002. By W. Nothdurft. Random House, Toronto. 242pp., illus. U.S. $24.95, CAD $37.95. *+Mammals of North America. 2002. By R. W. Kays and D. E. Wilson. Princeton University Press, Princeton. 240 pp., illus. Cloth U.S. $49.50: paper U.S. $19.95. +The mountain white-crowned sparrow: migration and reproduction at high altitude. 2002. By M. L. Morton. Studies in Avian Biology No. 24. Cooper Ornithological Society, Camarillo, California. 236 pp., illus. U.S. $27. *North American owls: biology and natural history. 2002. By P. A. Johnsgard. 2" edition. xiii + 298 pp., illus. + plates. U.S. $49.95. Botany Alpine plants of North America. 2002. By G. Nichols, Timber Press, Portland, Oregon. c 352 pp., illus. ¢ U.S. $49.95. A cactus odyssey: journeys in the wilds of Bolivia, 352 Peru, and Argentina. 2002. By J. D. Mauseth, R. Kiesling, and C. Ostolaza. Timber Press, Portland, Oregon. 306 pp., illus. U.S. $39.95. Carnivorous plants of the United States and Canada. 2002. By D. Schnell. 2" edition. Timber Press, Portland, Oregon. 468 pp., illus. U.S. $39.95. The cycads. 2002. By L. M. Whitelock. Timber Press, Portland, Oregon. 532 pp., illus. U.S.$59.95. +The ecology of plants. 2002. By J. Gurevitch, S. M. Scheiner, and G. A. Fox. Sinauer Associates, Sunderland, Massachusetts. xvi + 532 pp., illus. U.S. $89.95. *Flora of Glacier National Park, Montana. 2002. By P. Lesica. Oregon State University Press, Covallis. 512 pp., illus. U.S. $32.95. The illustrated encyclopedia of trees. 2002. By J. White. Timber Press, Portland Oregon. c800 pp., illus. cU.S. $79.95. *+Lewis Clark’s field guide to wild flowers of field and slope in the Pacific northwest. 2002. By L. J. Clark. Edited by J. Trelawny. 3™ edition. Harbour Publishing, Madeira Park, British Columbia. 80 pp., illus. $9.95. Portraits of Himalayan flowers. 2002. By T. Yoshida. Timber Press, Portland, Oregon. c120 pp., illus. U.S. $39.95, *Trees, shrubs, and vines for attracting birds. 2002. By R. M. DeGraaf. 2"4 edition. University Press of New England, Lebanon, New Hampshire. xiii + 169 pp., illus. U.S. $19.95. Environment Anatomy of a conflict: identity, knowledge, and emo- tion in old growth forests. 2002. By T. Satterfiled. UBC Press, Vancouver. 224 pp., illus. $85. +The beachcomber’s guide to seashore life of California. 2002. By J. D. Sept. Harbour Publishing, Madeira Park, British Columbia. 312 pp., illus. $28.95. tConservation biology. 2002. By A.S. Pullin. Cambridge University Press, New York. xii + 345 pp., illus. Cloth U.S. $120; paper U.S. $45. Handbook of ecological restoration, volume 1: princi- ples of restoration. 2002. Edited by M. R. Perrow and A. J. Day. Cambridge University Press, New York. 432 pp., illus. U.S. $100: volume 2: restoration practice. 650 pp., illus. U.S. $100; 2 volume set U.S.'$175. Integrating landscape ecology into natural resource management. 2002. Edited by J. Liu and W. W. Taylor. Cambridge University Press, New York. 500 pp., illus. Cloth U.S. $120: paper U.S. $45. Introductory ecology. 2002. By P. Cotgreave and I. THE CANADIAN FIELD-NATURALIST Vol. 116 Forseth. Blackwell Publishing, Oxford United Kingdom. 288 pp., illus. ,21.99. Islands of the Arctic. 2002. By J. Dowdeswell and M. Hambrey. Cambridge University Press, New York. 320 pp., illus. U.S. $38. +Life, temperature, and the earth: the self-organizing biosphere. 2002. By D. Schwartzman. Columbia University Press, New York. 241 pp., illus. U.S. $27.50. *The last island: a naturalist’s sojourn on Triangle Island. 2002. By A. Watt. Harbour Publishing, Madeira Park, British Columbia. 192 pp., illus. $34.95. Plant animal interactions. 2002. Edited by C. M. Herrera and O. Pellmyr. Blackwell Publishing, Oxford, United Kingdom. 336 pp., illus. ,35. + Quantitative conservation biology: theory and practice of population viability analysis. 2002. By W. F. Morris and D. F. Doak. Sinauer Associates, Sunderland, Massa- chusetts. xvi + 480 pp., illus. U.S. $39.95. Restoration of the Great Lakes: promises, practices, and performances. 2002. By M. Sproule-Jones. UBC Press, Vancouver. 160 pp., illus. $75. +The sacred balance: a visual celebration of our place in nature. 2002. By D. Suzuki and A. McConnell. Greystone Books (Douglas and McIntyre), Vancouver. 151 pp., illus. $55. +Spatial optimization for managed ecosystems. 2002. By J. Hof and M. Bevers. Columbia University Press, New York. xii + 257 pp., illus. Cloth U.S. $69.50; paper U.S. $32.50. Teaching sustainability: towards curriculum greening. 2002. Edited by W. L. Filho. Peter Lang Scientific Publishers, Bern. 570 pp. Euro 69.80. *When the wild comes leaping up: personal encounters with nature. 2002. Edited by D. Suzuki. Greystone Book, Douglas and McIntyre, Vancouver. 240 pp. $32.95. Miscellaneous *The backyard astronomer’s guide. 2002. By T. Dickinson and A. Dyer. Firefly Books, Willowdale, Ontario. 320 pp., illus. $49.95. +The canoe: a living tradition. 2002. Edited by J. Jen- nings. Firefly Books, Willowdale, Ontario, 271 pp., illus. $59.95. The cost of climate policy. 2002. By M. Jaccard, J. Nyboer, and b. Sadownik. UBC Press, Vancouver. 288 pp., illus. Cloth $85; paper $29.95. *The emperor of nature: Charles-Lucien Bonaparte and his world. 2000. By P. T. Stroud. University of Pennsylvania Press, Philadelphia. 371 pp. U.S. $34.95. 2002 BOOK REVIEWS 353 +The hydrogen economy. 2002. By J. Rifkin. Tarcher/ Putnam, New York. 283 pp., U.S. $24.95. *John Keast Lord: materials for a life. 2002. By D. Baker. Backhuys Publishers, Leiden, Netherlands. 78 pp.., illus. Euro 22. *The power of place: Charles Darwin, the origin and after — the years of fame. 2002. By J. Browne. Alfred A Knopf, New York. 624 pp., illus. U.S. $37.50. Books for Young Naturalists Salmon. 2002. By D. Hodge. Kids Can Press, Tona- wanda, New York. 32 pp., illus. U.S. $10.95. Seashells by the seashore. 2002. By M. Berkes. Dawn, Nevada City, California. 32 pp., illus. U.S. $8.95. *Assigned for review +Available for review News and Comment Errata The Canadian Field-Naturalist 115(4) Baird, R. W. 2001. Status of Killer Whales, Orcinus orca, in Canada. Canadian Field-Naturalist 115(4): 676-701. Figure 4. This figure should have included two pho- tographs, one (left) showing the dorsal fin of resident Killer Errata The Canadian Field-Naturalist 116(1) Inside front cover, bottom, caption for front cover: “Wood Bison, Bison bison bison” should read “Bison, Bison bison’. The population originally native to Wood Buffalo Park was designated Wood Bison as Bison bison athabascae Rhodes, 1898, Proceedings of the Academy of Natural Science, Philadelphia 49: 498, but the taxomonic distictiveness has long been debated. If regarded as distinct, the Plains Bison is Bison bison bison (Linnaeus, 1758) [See RENEW: National Recovery Plan Number 21: The Wood Bison, Bison bison athabascae, October 2001, 50 pages, Recovery Secretariat c/o Canadian Wildlife Service, Ottawa, K1A 0H3, noted in Whales, and one (right) showing the dorsal fin of transient Killer Whales. Only one photograph, of transient Killer Whales, is included in the Figure. Resident Killer Whales are shown in Figure 3, on opposite page. News and Comment in The Canadian Field Naturalist 116(1): 178]. The date of the cover photograph 116(1) was 7 July 1951, and the location was about 62 miles from Fort Smith on the road to Pine Lake. These Bison were probably hybrids Bison bison bison X athabascae, or simply Bison bison. The Plains Bison had been introduced in the years 1925 to 1928 so they had more than 20 years to interbreed by 195i. [W. A. Fuller 13 December 2002]. On page 146, left column, line 46, 1923 should be 1823, and the reference is to Sabine, Zoological Appendix V in Franklin 1823. Recovery: An Endangered Species Newsletter (20) March 2002 Published by the Canadian Wildlife Service this issue contains: RECOVERY HIGHLIGHTS: Strategy drafted — Saving the martens — New Law [Newfoundland and Labrador]; PROFILE: Team’s efforts paying off [David Wylynko]; sPE- CIAL REPORT: Kids, space, and species; NEWS BITES: Reintroducing snails — Recovering the copper redhorse — Orchid research underway — Updates: COSEWIC, RENEW — Deerberry recovery — Plan drafted [whooping crane] — Coastal flora at risk — FIELD NOTES: Monitoring trout [Ed Snucins] — The importance of animal behaviour [Abby Schwartz] — Tracking toads [David Green]; ANNOUNCEMENTS: New publications — Upcoming events — Site seeing; FEATURED SPECIES: Conserving a rare plant [Alain Gouge]. Contact: Recovery, Canadian Wildlife Service, Environment Canada, Ottawa, Ontario K1A 0H3. Web site: www.cws-scf.ec.gc/recovery/archive.htmla Froglog: Newsletter of the Declining Amphibian Populations Task Force (50) Number 50, April 2002, Celebrating Froglog’s Golden Jubilee! CONTENTS: A Message from DAPTF’s Co-Founder (David Wake) — DAPTF Seed Grants (Tim Halliday, DAPTF International Director) — Breeding Pond Survey in Hungary: an Example of Successful Cooperation (Tibor Kovacs & Miklos Papp) — Green and Golden Bell Frogs on Kooragang Island [Australia] (Robert Browne Chairperson, Friends of the Hunter Bell Frogs) — Meeting in the Basque Country (Tim Halliday) — California-Nevada Working Group Meeting Report (David Bradford, Working Group Chair) — A Comparative Study of the Amphibian Com- munity at Three Sites in Penang State, Malaysia — Froglog Shorts —- Publications of Interest. Froglog is the bi-monthly newsletter of the Declining Amphibian Populations Task Force of The World Con- servation Union (IUCN)/Species Survival Commission (SSC) and is supported by The Open University, The World Congress of Herpetology, and Harvard University. The newsletter is Edited by John W. Wilkinson, Department of Biological Sciences, The Open University, Walton Hall, Milton Keynes, MK7 6AA, United Kingdom; e-mail: daptf@open.ac.uk. Funding for Froglog is underwritten by the Detroit Zoological Institute, P. O. Box 39, Michigan 48068-0039, USA. 354 2002 THE OTTAWA FIELD-NATURALISTS’ CLUB AWARDS 355 National Recovery Plan (22): Piping Plover March 2002 The National Recovery Plan for the Piping Plover (Charadrius melodus) by the Prairie and Atlantic Piping Plover Recovery teams (J. P. Goossen, D. L. Amirault, J. Arndt, R. Bjorge, S. Boates, J. Brazil, S. Brechtel, R. Chiasson, G. N. Corbett, R. Curley, M. Elderkin, S. P. Flemming, W. Harris, L. Heyens, D. Hjertaas, M. Huot, B. Johnson, R. Jones, W. Koonz, P. Laporte, D. McAskill, R. I. G. Morrison, S. Richard, F. Shaffer, C. Stewart, L. Swanson, and E. Wiltse) is now available as Number 22, 47 pages, from Recovery Secretariat c/o Canadian Wildlife Service, Environment Canada, Ottawa, Ontario K1A OH3, Canada; e-mail RENEW-RESCAPE @ec.gc.ca; See also RECOVERY web site: English: http://www.speciesatrisk.gc.ca/sar/efforts/index.htm Francais: http://www.especesenperil.gc.ca/eep/efforts/index.htm Point Pelee Natural History News 2(1) Spring 2002 This newsletter for Point Pelee National Park, Ontario, is edited by Alan Wormington (e-mail: wormington@juno. com). Editorial Assistants are M. Lea Martell and Matthew J. Smith. The web site is www.wincom.net/~fopp/ Natural_History_News.htm ARTICLES: The Point Pelee Lesser Nighthawk: A Unique Record for Ontario and Canada (Alan Wormington) — Noteworthy bird records: December 2001 to February 2002 (Alan Wormington) — “Saskatchewan” Horned Lark: New to Ontario (Alan Wormington) — Point Pelee Christmas Marine Turtle Newsletter (96) The April 2002 issue, 40 pages, contains: EDITORIAL: Hype (Nicholas Mrosovsky) — ARTICLES: Marine turtle nesting in Kuriat Islands, Tunisia, 2000 — Observations on the first day dispersal of neonatal Hawksbill Turtles (Eretmochelys imbricata) — In-water survey of Hawksbill Turtles at Kuna Yala, Panama — Evidence for Leatherback Sea Turtle (Dermochelys coriacea) nesting in Arraial do Cabo, State of Rio de Janeiro, and a review of occasional Leatherback nests in Brazil — Sea Turtles nesting in the South Bay of Great Nicobar Island — Marine turtles nesting at St. Martin’s Island, Bangladesh. Notes: A record of the Leatherback Sea Turtle (Dermo- chelys coriacea) from Cambodia — LETTER TO THE EDI- TORS: Viewpoint: Sea turtle management in Bangladesh Bird Count: December 17, 2001 (Sarah Rupert) — Point Pelee Butterflies: Annual Summary for 2001 (Alan Worm- ington) — Upcoming EVENTS AND OUTINGS. Subscription rates are Canada: CAN $15 (one year) or $30 (two years); International: US $15 (one year) or $30 (two years). Send payment (and e-mail address, optional) to The Friends of Point Pelee, 1118 Point Pelee Drive, Leamington, Ontario N8H 3V4 Canada. Issues are mailed in March, June, September, and December, and back issues are avail- able for $15 per Volume/ $5 per issue (postage paid). — ANNOUNCEMENTS — NEWS & LEGAL BRIEFS — RECENT PUBLICATIONS. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Research Group, School of Biological Sciences, University of Wales, Swansea, SA2 8PP Wales, United Kingdom; e-mail MTN @swan.ac.uk; Fax +44 1792 295447. Subscriptions to the MTN and donations towards the production of MTN and its Spanish edition NTM [Noticiero de Tortugas Marinas] should be sent to Marine Turtle Newsletter c/o Chelonian Research Foundation, 168 Goodrich Street, Lunenburg, Massachusetts 01462 USA; e-mail RhodinCRF @aol.com; fax + 1 978 582 6279. MTN website is: The Boreal Dip Net: Newsletter of the Canadian Amphibian and Reptile Conservation Network 6(2) April 2002 Contents: Editor’s note [Kerrie Serben]; Charlottetown West: Revisiting the PEI Meeting at Eastern Ontario Biodiversity Museum 27 January 2002 [Aleta Karstad] — Projects en herpetologie au Quebec [Jacques Jutras et col- laborateurs] — Ontario snake and lizard advisory group formed [David Seburn] — Herpwatch [Aquatics, Banff National Park Warden Office] — Considerations of the inadequacy of wildlife reserve sizes in Ontario for the con- servation of reptiles and amphibians (Shane R. de Solla] — Habitat use and movement patterns of Northern Alligator Lizards and Western Skinks in southeastern British Columbia [Pamela Rutherford and Patrick Gregory] — Don’t be caught out standing in a pond the spring! [FrogWatch, NatureWatch Program, joint venture of Canadian Nature Federation and the Ecological Monitoring and Assessment Network Coordinating Office, Environment Canada] — Field notes: Northern Dusky Salamander [Glenn Barrett and Kim Smith] — Message in a bottle: A new beer launched in support of species at risk [Andree Gendron] — The new atlas of the reptiles and amphibians of Vermont is now available [James S. Andrews] — Herpetological activi- ties in eastern Canada in 2002: monitoring-conservation- education-research [Martin Ouellet]. For information on membership in the Canadian Am- phibian and Reptile conservation society ($10 students, $16 non-students] contact Bruce Pauli, Canadian Wildlife Service, National Wildlife Research Centre, 100 Gamelin Boulevard, Hull, Quebec K1A OH3. Web site: http://www.carenet.ca/ 356 THE CANADIAN FIELD-NATURALIST Vol. 116 Alberta Wildlife Status Reports numbers 40, 41, 42, 43, 44 The Fisheries and Wildlife Management Division of the 42. Status of the Prairie Falcon (Falcon mexicanus) in Alberta Natural Resource Status and Assessment Branch, Alberta, by Dale Paton. 28 pages. Alberta Environmental Protection, has released new 43. Status of the American Badger (Taxidea taxus) in Wildlife Status Reports. The Series Editors for reports 40 to Alberta, by Dave Scobie. 17 pages. 44 are Sherry Frazer and Robin Gutsell and the illustrations 44. Status of the Yuca Moth (Tegeticula yuccasella) in for all five are by Brian Huffman. For a listing earlier num- Alberta, by Donna Hurlburt. 21 pages. bers in the series, see The Canadian Field-Naturalist For copies contact the Information Centre - Publications 112(1): 169 for 1-11; 113(2): 311 for 12-17; 113(4): 686 Alberta Environment/Alberta Sustainable Resource for 18-21; 114(1): 151 for 22-25; 115(2): 390 for 26-31; | Development, Fish and Wildlife Division, Main Floor, 115(3): 000 for 32-36, 116(1): 000-000. Recent reports Great West Life Building, 9920 - 108 Street, Edmonton, issued in 2002 are 40, 41, 42 and 44, dated March 2002: 43 Alberta TSK 2M4 Canada [telephone: (780) 422-2079], OR is dated May 2002): Information Service Alberta Environment/Sustainable 40. Status of the Banff Springs Snail (Physella johnsoni)in Resource Development, #100, 3115 - 12 Street NE, Alberta, by Dwayne A. W. Lepitzki. 29 pages. Calgary, Alberta T2E 7J2, Canada [telephone: (403) 297- 41. Status of the Shortjaw Cisco (Coregonus zenithicus) in 3362]; web site: http://www3.gov.ab.ca/srd/fw/status/ Alberta, by Mark Steinhilber. 23 pages. index.html. Editor’s Report for Volume 115 (2001) Mailing dates for issues in volume 115 were: (1) | Chow and the pre-press staff for galleys and correc- 10 July 2001; (2) 14 December 2001; (3) 30 April tions as well as those in the printing and bindary sec- 2002; (4) 9 August 2002. A summary of membership _ tions whose efforts make each issue possible. Wanda and subscriber totals 2001 is given in Table 1. The J. Cook proof-read the galleys and Bill Cody as number of articles and notes in volume 115 is sum- Business Manager handled all reprint requests and marized in Table 2 by topic; totals for Book Reviews _ bills and oversaw and proofed the compilation of the and New Titles are given in Table 3, and the distri- Index prepared by Leslie Durocher. Wilson Eedy con- bution of content by page totals per issue in Table 4. _ tinued as Book-Review Editor (his report follows). Council continued making 80% of the annual inter- Manuscripts (excluding book reviews, notices, and est from the Manning Fund and other capital avail- _ teports) submitted to The Canadian Field-Naturalist totalled able to The Canadian Field-Naturalist to offset the 125 in 2001, down 29 from 154 in 2000. The following publication cost of northern papers where authors reviewed papers submitted in late in 2000 and in 2001: and institutional contributions were insufficient to Associate’ Eaitors” Ro idersot 2 ee Nature (3); W. B. Ballard, Texas Tech University, Lubbock cover page charges. f (10); C. D. Bird, Erskine, Alberta (26); B. W. Coad, St. Joseph Print Group, Ottawa (now including the — Canadian Museum of Nature (9); R. Campbell, Ottawa, (6); former M.O.M. Printers) set and printed the journal. P.M. Catling, Agriculture and Agri-food Canada, Ottawa Special thanks are due Emile Holst, and to Cecilia (14); A. J. Erskine, Canadian Wildlife Service, Sackville, TABLE |. The 2001 circulation of The Canadian Field-Naturalist (2000 in parenthesis). Membership totals from Annual Report of the Ottawa Field-Naturalists’ Club, January 2002; subscription totals compiled by W. J. Cody. Forty percent of membership dues and 100% of subscriptions go to publication of The Canadian Field-Naturalist. Members vote on Club affairs, subscribers and institutions do not. Canada USA Other Totals Memberships Family & individual 941 (910) 30 (32) 7 (6) 978 (948) Subscriptions Individuals 189 (184) 63 (61) 3 (6) 255 (251) Institutions 174 (178) 262 (255) 38 (38) 474 (471) Totals 363 (362) $25 (316) 41 (44) 729 (722) TOTALS 1304 (1272) 320 (348) 48 (50) 1707 (1670) Note: 22 countries are included under “Other” (outside Canada and United States): Austria, Belgium, Brazil, Denmark (2), United Kingdom (9: including | to Scotland), Finland (2), France (3: including | to St. Pierre & Miquelon), Germany (2), Iceland, Ireland, Japan, Mexico, Netherlands (3), New Zealand, Norway (4), Poland, Russia, South Africa, Spain (3), Sweden (2), Switzerland (2), Trinidad and Tobago. 2002 THE OTTAWA FIELD-NATURALISTS’ CLUB AWARDS 357 TABLE 2. Number of articles and notes published in The Canadian Field-Naturalist Volume 115 (2001) by major field of study. Subject Articles Notes Total Mammals 19 8 27 Birds 12 9 21 Amphibians + reptiles 3 3 6 Fish 15 1 16 Invertebrates 3 0 5 Plants 14 4 18 Tributes 2 0 2 Totals 70 25 95 New Brunswick (31); W. O. Pruitt, Jr., University of Manitoba, Winnipeg (55). Other reviewers: R. Alvo, Gatineau, Quebec; R. Anderson, Seattle Aquarium, Washington; M.N. Arai, Pacific Biological Station, Nanaimo, British Columbia; A. Bailey, University of Alberta, Edmonton, Alberta; W. M. R. Barclay, University of Calgary, Alberta (2); W. Berg, Minnesota Department of Natural Resources, Grand Rapids; J. R. Bider, Ecomuseum, Ste-Anne-de-Bellevue, Quebec (4); M. Blouw, University of Northern BC, Prince George; J. Bowman, Carleton University, Ottawa, Ontario; C. Braun, Colorado State University, Fort Collins (2); S. Breck, Colorado State University, Fort Collins; A. Brinckmann-Voss, Sooke, British Columbia; I. Brodo, Canadian Museum of Nature, Ottawa, Ontario; R. J. Brooks, University of Guelph, Ontario (3); T. Brunjes, Texas Technical University, Lubbock, Texas; M. Cadman, Canadian Wildlife Service, Guelph, Ontario; D. Colwell, Canada Agriculture and Agri-Food, Lethbridge; L. Carbyn, CWS, Edmonton, Alberta (4); J. G. Casey, Narragansett, Rhode Island; J. Cayouette, Agriculture and Agri-food Canada, Ottawa; P. Cochran, Saint Mary’s University, Winona, Minnesota; W. J. Cody, Agriculture and Agri-food Canada, Ottawa, Ontario (8); J. A. Cook, Museum, University of Alaska, Fairbanks; V. Crichton, Manitoba Natural Resources, Winnipeg; E. J. Crossman, Royal Ontario Museum, Toronto (4); D. Cuddy, OMNR, Kemptville, Ontario; S. J. Darbyshire, Agriculture and Agri- food Canada, Ottawa (2); R. W. Davies, University of Calgary, Alberta; M. W. Demarchi, LGL Limited, Sidney, British Columbia; K. De Smet, Manitoba Conservation, Meleta; A. Dextrase, Ontario Ministry of Natural Resources, Peterborough; G. W. Douglas, BC Data Centre, Victoria (2); J. Dubois, Manitoba Natural Resources, Winnipeg (3); B. A. Ford, University of Manitoba, Winnipeg; R. Fortin, TABLE 3. Number of reviews and new titles published in Book Review section of The Canadian Field-Naturalist Volume 114 by topic. Reviews New Titles Zoology 16 112 Botany 9 24 Environment 11 61 Miscellaneous 3 18 Young Naturalists = 56 Totals if / ZV TABLE 4. Number of pages per section published in The Canadian Field-Naturalist Volume 115 (2001) by issue. (1)i, Qi) + Cie ts Vata Articles 1G} AAD 104 0 £594? S72 Notes Ly 16 18 0 51 Book Reviews 12 9 14 26 61 Tributes 24 3 | CFN/OFNC Reports” 3 / 4 14 News and Comment 2 i 2 2 8 Index 35 35 OFNC Publications 1 1 Advice to Contributors 1 1 Totals 198 196. 148, 228 770 “Total pages for book review section include both reviews and new titles listings. “Includes CFN Editors reports (issue 2), OFNC Annual Business Meeting (3) and OFNC Awards (4). Université du Québec 4 Montréal, Québec (2); B. Freedman, Dalhousie University, Halifax, Nova Scotia; A. J. Gaston, Canadian Wildlife Service, Gatineau, Quebec (2); L. Gillespie, Canadian Museum of Nature, Ottawa; P. T. Gregory, University of Victoria, British Columbia (2); I. Hatter, British Columbia Ministry of Environment, Lands and Parks, Victoria; D. Heard, British Columbia Ministry of Environment, Lands and Parks, Prince George; B. J. Hearn, Natural Resources Canada, Corner Brook, Newfoundland: D. Helm, University of Alaska, Palmer; J.D. Henry, Klune National Park, Yukon; L. V. Hills, Cochrane, Alberta; E. Holm, Royal Ontario Museum, Toronto; G. Holroyd, Canadian Wildlife Service, Edmonton, New Brunswick: C. S. Houston, Saskatoon, Saskatchewan (3); P. Hurley, Department of Fisheries and Oceans, Darmouth, Nova Scotia; T. C. Hutchinson, Trent University, Peterbourgh, Ontario; P. James, Royal Saskatchewan Museum, Regina; R. James, Sutherland, Ontario (3); H. M. Jahns, Heinrich Heine Universitat, Dusseldorf, Gemany; C. E. Kay, Utah State University, Logan; P. Keddy, Carleton Place, Ontario (3); M. R. Lein, University of Calgary, Calgary; L. E. Licht, York University, North York; J. W. Lish, Oklahoma State University, Stillwater; R. MacCulloch, Royal Ontario Museum, Toronto; C. Machtans, Candian Wildlife Service, Yellowknife, Northwest Territories; J. Madill, Canadian Museum of Nature, Ottawa; F. F. Mallory, Laurentian University, Sudbury (2); A. Martel, Canadian Museum of Nature, Ottawa (2); J. Mather, University of Lethbridge, Alberta; T. O. Matson, Cleveland Museum of Natural History, Ohio; J. Maunder, Newfoundland Museum, St. John’s (2); D. F. McAlpine, New Brunswick Museum, Saint John (4); R. McCaffrey, Texas Techical Unversity, Lubbock (2); B. McCune, Oregon State University, Corvalis; W. B. McGillivray, Alberta Provincial Museum, Edmonton (3); L. David Mech, Raptor Center, University of Minnesota, St. Paul (2); J. S. Millar, University of Western Ontario, London; J. Metcalfe-Smith, National Water Research Institute, Burlington, Ontario; G. Morrison, Canadian Wildlife Service, Gatineu, Quebec; K. Murphy, Yellowstone Park, Wyoming; D. W. Nagorsen, Victoria, British Columbia (7); R. W. Nero, Manitoba Natural Resources, Winnipeg, Manitoba; W. A. C. Nixon, Canadian Wildlife Service, Yukon; L. Noel, LGL Alsaka Research Associates, OOOO eee 358 THE CANADIAN FIELD-NATURALIST Anchorage; M. J. Oldham, OMNR, Peterborough (2); S. Petrie, Bird Studies Canada, Port Rowan, Ontario (2); R. S. Phillips, Texas Technical University, Lubbock; G. Proulx, Alpha Research and Management, Sherwood Park, Alberta (4); J. C. Ray, University of Toronto, Ontario; G. Redmond, New Brunswick Department of Natural Resouces, Fredericton; J. Reed, Texas Technical Universtiy, Lubbock; R. Reeves, Okapi Wildlife Associates, Hudson, Quebec (2); T. E. Reimchen, University of Victoria, British Columbia; M. Robert, Canadian Wildlife Service, Sainte Foy, Quebec; G. J. Robertson, CWS, Mount Pearl, Newfoundland; R. C. Rosatte, Ontario Ministry of Natural Resourses, Trent University, Peterborough; W. M. Samuel, University of Alberta, Edmonton; J.-P. Savard, Canadian Wildlife Service, Ste. Foy, Quebec; F. Scott, Acadia University, Wolfville, Nova Scotia (4); D. Scheel, Alaska Pacific University, Anchorage; W. B. Schofield, University of British Columbia, Vancouver; M. Schroeder, Washington Department of Fish and Wildlife, Bridgeport; F. W. Schueler, Eastern Ontario Biological Museum, Kemptville, Ontario (5); B. Scott, Kingston, Ontario (2); S.G. Sealy, U Manitoba, Winnipeg, Manitoba (2); B. Skomal, Martha’s Vinyard Research Station, Vinyard Haven, Massachusetts; C. M. M. Smits, Department of Renewable Resources, Yukon Territory; R. T. Stevens, University of Memphis, Tennessee; K. W. Stewart, University of Manitoba, Winnipeg (2); I. Stirling, Canadian Wildlife Service, Edmonton, Alberta (3); J. Theberge, Vol. 116 University of Waterloo, Ontario; R. Thiel, Sandhill Wildlife Area, Babcock, Wisconsin; I. Thompson, Canadian Forest Service, Sault Ste. Marie, Ontario (6); T. N. Todd, United States Geological Survey, Ann Arbor, Michigan; N. Ver- beek, Simon Fraser University, Burnaby, British Columbia; A.M. Veitch, Northwest Territories Department of Resources, Wildlife and Economic Development, Norman Wells; D. Vitt, University of Alberta, Edmonton; K. Wiebe, University of Saskatchewan, Saskatoon; E. Wiltse, Saskatchwan Environment and Resource Management, Regina; K. K. Wright, Cornell College, Mount Vernon, Iowa; A. Wydeven, Wisconsin Department Natural Resouces, Park Falls. I am also indebted to Eleanor Zurbrigg, President of the Ottawa Field-Naturalists’ Club and the Club Council for continuing support of the journal; Chairman Ron Bedford and the Publications Committee of the OFNC for editorial encouragement and support, to the Canadian Museum of Nature for access to its library and the facilities at the Natural Heritage Building, 1740 Pink Road, Aylmer, Quebec, and to Joyce for everything else. FRANCIS R. COOK Editor Book-review Editor’s Report for Volume 115 (2001) This has been a very eventful year for your book- review editor and I want to make sure everyone knows where I am. After 28 years in one location, I have moved my address. From one remote, dead- end, ANSI and ESA area to a little less remote but still green-belt location, about 9 ha on the edge of the Mad River, just southwest of Barrie. Reviewers will have to forgive any tardiness due to files still in boxes, my still working in Africa, and major renova- tions at home. These latter include one huge frog pond and bird habitat creation in the front lawn. For those who want to find me: RR# 1, Glencairn, Ontario LOM 1KO; e-mail terfa@ geoniger.com. Book reviewing for the Canadian Field-Naturalist in 2001 was a little slower than usual. Although we received a large number of complimentary titles from publishers (102), only 65 were sent to review- ers and 56 reviews published in the journal. The number of new titles listed, 261, was also low, but partly due to only listings in 3 issues. This means we have a large number of books still available for reviewers and welcome contacts from both old or new reviewers. Just let me know the topics that you are interested in and I will send suggestions that you can pick from. You can also suggest titles from those listed as available in New Titles or even suggest rel- evant new books which I have missed. Most publish- ers will provide complimentary copies on request. The book becomes the property of the reviewer and the review is published in our journal. WILSON EEDY Book-review Editor oreo TABLE OF CONTENTS (concluded) MISCELLANEOUS: Finding Order in Nature: The Naturalist Tradition from Linnaeus to E. O. Wilson — Excavation . New Titles News and Comment The Canadian Field-Naturalist Errata: 115(4) — Errata: 116(1) — Recovery: An Endangered Species Newsletter (20) March 2002 — Froglog: Newsletter of the Declining Amphibian Populations Task Force (50) — National Recovery Plan (22): Piping Plover March 2002 — Point Pelee Natural History News 2(1) Spring 2002 — Marine Turtle Newsletter (96) — The Boreal Dip Net: Newsletter of the Canadian Amphibian and Reptile Conservation Network 6(2) April 2002 — Alberta Wildlife Status Reports numbers 40, 41, 42, 43, 44. Editor’s Report for Volume 115 (2001) Book Review Editor’s Report for Volume 115 (2001) Mailing date of the previous issue 116(1): 30 November 2002 349 351 354 356 358 THE CANADIAN FIELD-NATURALIST Volume 116, Number 2 Articles Winter habitat selection at three spatial scales by American Elk, Cervus elaphus, in west-central Alberta PAUL F. JONES and ROBERT J. HUDSON Notable vascular plants from Alaska in Wrangell-St. Elias National Park and Preserve with comments on the floristics Mary B. Cook and CARL A. ROLAND Notes Black Bear, Ursus americanus, hair and Apple Trees, Malus pumila, in northeast North America DAVID H. HIRTH, JOSEPH M. A. PETTY, and C. WILLIAM KILPATRICK Large clutch size of a Burrowing Owl, Athene cunicularia, found in Saskatchewan L. DANIELLE TODD and JOANN SKILNICK Dispersal by a male American Marten, Martes americana DorotuHy M. FECSKE and JONATHAN A. JENKS Territorial marking by lone male Gray Wolves, Canis lupus RONALD N. SCHULTZ and PAMELA C. WILSON Estimating the weight of Wolves, Canis lupus, from chest girth measurements JOHN P. HART, and DAvID H. JAMIESON Differential use of a Wolf, Canis lupus, pack territory edge and core L. DAviD MECH and ELIZABETH K. HARPER Extension de l’aire de distribution de la Salamandre a quatre doigts, Hemidactylium scutatum, dans |’ est du Québec, et notes sur |’ habitat JEAN-FRANCOIS DESROCHES et BENOIT COUTURE Reproductive characteristics of Northern Pocket Supe. Thomomys talpoides, in Alberta alfalfa fields GILBERT PROUBX Evidence of a second litter in Northern Pocket Gophers, Thomomys talpoides GILBERT PROULX and PAMELA J. COLE Narwhal, Monodon monoceros, near western Baffin Island, Nunavut, Canada MarkK L. MALLORY and ANDREW B. DIDIUK Tributes A tribute to Edward Warren Greenwood (1918-2002), Canadian Orchidologist Joyce M. REDDOCH and ALLAN H. REDDOCH Book Reviews Zoology: Butterflies of British Columbia (including Western Alberta, Southern Yukon, the Alaska Dynamics of the Boreal Forest: The Kluane Project — The Canadian Rockies Guide to Wildlife Watching: The Best Places to See and Appreciate Animals in Their Natural Habitat — Four Wings and a Prayer — Radio Tracking and Animal Populations — The Birds of Ecuador: Status, Distribution and Taxonomy (Volume I), Field Guide (Volume IJ) — Sylvia Warblers — Bats of Papua New Guinea — A Field Guide to Birds of Peru — Birds of Delaware — Checklist of the Birds of Northern South America — Birds of Southern South America and Antarctica — Whales and Other Marine Mammals of British Columbia and Alaska — Guia de las Aves de Espafia Botany: Magical Mushrooms, Mischievous Molds — Spatial Pattern Analysis in Plant Ecology — Rare Vascular Plants of Alberta Environment: The Global Environment in the Twenty-first Century: Prospects for International Cooperation — Reading the Entrails: An Alberta Ecohistory — Spacial Optimization for Managed Ecosystems — The Amber Forest — The Natural History of an Arctic Oil Field, Development and the Biota ISSN 0008-3550 3 9088 01226 6144 The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Bee ed on | = = fas | _ y E y _ = y 7 Zz = = Volume 116, Number 3 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons ; Her Excellency The Right Honourable Adrienne Clarkson, C.C., C.M.M., C.D. Governor General of Canada His Excellency John Ralston Saul, C.C. 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 infor- mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain- ing or restoring environments of high quality for living things. Honorary Members Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope Donald M. Britton R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. Irwin M. Brodo Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch William J. Cody John M. Gillett Theodore Mosquin Mary E. Stuart Francis R. Cook C. Stuart Houston Eugene G. Munroe John B. Theberge Ellaine Dickson George F. Ledingham Robert W. Nero Sheila Thomson 2002 Council ‘ President: Eleanor Zurbrigg Vice-President: Gary McNulty Ronald E. Bedford Francis R. Cook David W. Moore Roy John Rosanne Bishop Barbara Gaertner Robert Roach ; . Irwin Brodo Diane Lepage Stanley Rosenbaum Recording Secretary: Ken Allison John Cameron Diane Holms Louise Schwartz Treasurer: Frank Pope William J. Cody David Hobden David Smythe Kathy Conlan Beverly McBride Dorothy Whyte To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, Box P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net. For information on Club activities telephone (613) 722-3050 or check http//www.achilles.net/ofnc/index.htm The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. We acknowledge the financial support of the Government of Canada toward our mailing cost through the Publication Assistance Program (PAP), Heritage number 09477. Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1RO0; (613) 269-3211; e-mail: feook @achilles.net Copy Editor: Elizabeth Morton Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 759-1374 Book Review Editor: Dr. J. Wilson Eedy, 9074-32 Side Road, R.R.1, Glencairn, Ontario LOM KOC Canada e-mail: terfa@ geoniger.com Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen Charles D. Bird Brian W. Coad Donald F. McAlpine Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr. Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication should be addressed to the Editor and sent by postal mail. Exception: book reviews should go directly to Book Review Editor. Subscriptions and Membership Subscription rates for individuals are $28 per calendar year. Libraries and other institutions may subscribe at the rate of | $45 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $28 (individual) $30 (family) $50 | (sustaining) and $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and mem | bers (including USA) must add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, cover the Ottawa District and Local Club events. It is mailed to Ottawa area members, and available to those outside Ottawa o1 request. It is available to Libraries at $28 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate | P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: July-September 2002. (March 2003). ~ Cover: A mat of the Eastern Mosquito Fern, Azolla caroliniana, stranded on the mud substrate with other aquatic vegetation | and fallen leaves at Brown’s Inlet, Ottawa, after the draw-down of water in the Rideau Canal, October 1997. Photographed by S. J. Darbyshine (see pages 441-445). The Canadian Field-Naturalist Volume 116, Number 3 July—September 2002 Double-crested Cormorants, Phalacrocorax auritus, at Inland Lakes North of Lake Huron, Ontario ROBERT ALVO!, CHRIS BLOMME2 and D. VAUGHN WESELOH? ‘GAVIA Biological Services, 58, rue des Parulines, Hull, Québec JOA 1Z2 Canada *Department of Biology, Laurentian University, Sudbury, Ontario P3E 2C6 Canada 3Canadian Wildlife Service, Environment Canada, 4905 Dufferin Street, Downsview, Ontario M3H 5T4 Canada Alvo, Robert, Chris Blomme, and D. Vaughn Weseloh. 2002. Double-crested Cormorants, Phalacrocorax auritus, at inland lakes north of Lake Huron, Ontario. Canadian Field-Naturalist 116(3): 359-365. Three long-term data sets were used to document the use of inland lakes north of Lake Huron by Double-crested Cormorants (Phalacrocorax auritus). After the first reported sighting in the Sudbury Region in 1971, the species was observed regularly from 1979 onwards. It was not a summer resident in the area until 1987 and noteworthy multiple num- bers were not reported until 1991. These non-breeding birds may have originated as spring migratory “overshoots” from Georgian Bay, Lake Huron; birds observed in the autumn were probably migratory stragglers from farther north. It is unlikely that they came from the east or west because present day colonies there were not active yet. The only two known breeding sites in the region are at Kelley Lake and Lake Wanapitei. Breeding at Kelley Lake began in 1996; in Lake Wanapitei, breeding was confirmed in 1999, but may have begun earlier. These sightings represent the earliest movements of cormorants away from the Great Lakes since their resurgence there, and these birds may establish breeding sites at small inland lakes in this area. These movements into the Sudbury Region are not to be confused with daily foraging flights to inland lakes which are seen within several kilometres of the Great Lakes. Further research is required to study what impact cormorants might have on inland lake ecosystems and to determine whether cormorants are seen with increased frequency elsewhere in the Great Lakes watershed. Key Words: Double-crested Cormorant, Phalacrocorax auritus, colony, Sudbury, North Bay, Manitoulin, Ontario, expansion. The Double-crested Cormorant (Phalacrocorax auritus) has been undergoing a continent-wide popu- lation expansion since the mid- to late-1970s (Weseloh et al. 1979; Scharf and Shugart 1981; Hatch 1984). Nowhere has this been more dramatic than on the Great Lakes (Weseloh et al. 1995). This has caused widespread concern as cormorants are perceived as having severe negative impacts on at least three components of the ecosystem: (1) fish- eries (e.g., Smallmouth Bass, Micropterus dolomieu, in eastern Lake Ontario: Schneider et al. 1998); (2) vegetation (e.g., islands in the St. Lawrence River: Alvo 1996; and western Lake Erie: DVW, personal observations); and (3) other colonial species with which cormorants nest (e.g., Great Blue Herons, Ardea herodias, and Black-crowned Night-Herons, Nycticorax nycticorax: Wires and Cuthbert 1999*; * Jarvie et al. 1997). : In the early 1900s, the Double-crested Cormorant _ bred in the Gulf of St. Lawrence and on lakes in the *See Documents Cited section prairie provinces, and was “a rather uncommon though perhaps regular migrant on the Great Lakes” (Taverner 1919). The breeding population on the Great Lakes increased slowly from the first known colonization in 1913-1920 to around 900 pairs by the 1950s. However, reproductive failure caused by DDE-induced eggshell thinning and subsequent egg breakage reduced the population to less than 150 pairs by 1972; the species abandoned all breeding sites in Lake Michigan, Lake Superior, and the main body of Lake Huron (Postupalsky 1978*). Since then, the cormorant population has increased dra- matically, probably due to a combination of factors: reduced human persecution, declining levels of organochlorine contaminants (Ryckman et al. 1998), and an abundance of forage-base fish, notably Alewife (Alosa pseud oharengus) and Rainbow Smelt (Osmerus mordax) (Christie 1974). This abun- dance of forage-base fish followed the dramatic decline of Great Lakes stocks of predatory fish between the 1940s and 1960s, which has been attributed to predation by Sea Lampreys (Petromyzon marinus) and human over-exploitation (Christie 1974; Christie et al. 1987). 359 360 From 1973 to 1993, the breeding cormorant popu- lation on the Great Lakes increased over 300-fold to more than 38 000 pairs (Weseloh and Collier 1995). In 1991, about 40% of the Canadian Great Lakes cormorant population bred in Lake Huron, where the nesting population grew from nine colonies with 335 nests in 1980 to 41 colonies with 6018 nests in 1989, representing a mean annual growth rate of 38% (Blokpoel and Tessier 1997). In 2000, cormorants bred on numerous islands in Lake Huron, with most of the population concentrated along its north shore; i.e., off Manitoulin Island, in the North Channel, and along the north and east coasts of Georgian Bay (Blokpoel and Tessier 1997; D. V. Weseloh, unpub- lished data). In Ontario and the U.S. states bordering the Great Lakes, breeding cormorants are most numerous along the Great Lakes shoreline and offshore islands. In these situations where cormorant activity seems confined to the Great Lakes proper, a feeling of hav- ing “isolated” any perceived cormorant problem from inland habitats often seems to prevail. However, great fear has often been expressed by wildlife managers that, if cormorants began using small inland lakes, the perceived problems would escalate. This subject has received little published scrutiny. Here we use three long-term data sets to document the initial stages of expansion into small inland lakes north of Georgian Bay, Lake Huron. Methods This study is based on three data sets: (A), (B), and (C). (A) As part of a long-term loon study in the region of Sudbury, Ontario (Alvo et al. 1988), RA recorded all observations of aquatic birds. Obser- vations were made annually from 1982 to 1990, in 1993, and from 1996 to 1998. In 1982, 84 lakes were visited from mid-June to late-August. In 1996 the same 84 lakes, plus 10 added in 1988 (see below), were visited from mid-June to late October. In the other years a subset of 30 lakes (the same 30 every year) was visited from late August to mid- September. In addition, in 1988, 10 lakes were added to the study, and they were surveyed from late August to mid-September in succeeding years with observations as described above. Usually only one visit per lake was made each year, but in 1982 and 1996 two visits per year were made. Thus, the num- ber of lakes surveyed per year was as follows: 1982: 84; 1983-1987: 30; 1988-1990 and 1993: 40; 1996: 94; 1997-1998: 40. In total, 94 lakes were visited 746 times. They were located 35-135 km from the 381-m tall Inco smelter “superstack” near Sudbury. This study region, henceforth referred to as the “out- lying Sudbury region”, forms a large quadrilateral block, with its axis in a north-south direction, north of Georgian Bay (Figure 1). THE CANADIAN FIELD-NATURALIST Vol-Ll6 (B) The second data set was compiled by CB, who searched for cormorant observations in the Sudbury Region in the Manitoulin and Sudbury District Bird Reports [the monthly report of birders who subsequently formed the Sudbury Ornithology Society — SOS] for the 1972-1999 period (editors John Lemon and John Nicholson). These reports Summarized members’ observations made during each month. In 1972-1976, all monthly reports were based on the observations of at least six observers. In 1977-1998, monthly reports were based on observations of an average of 10.4 observers. For each bird species, historical or first time sightings were noted, as were the earliest spring and latest autumn observations and the range of summer resi- dency dates. The geographical area covered includes a roughly 40 km radius around the city of Sudbury; i.e., an area overlapping by about 40% with A (Figure 1). Observations by CB for the same 28- year period, and reports obtained during conversa- tions with local birders, were also included in this data set. For simplicity, this data set is referred to as the “SOS” data. (C) The third data set comprises 97 counts of cor- morants on Kelley Lake (339 ha), located 3 km southwest of the city (Sudbury) core. The counts were made by CB from 1972 to 1999 at various times during the breeding seasons (Figure 1). Kelley Lake is noted for its waterfowl migrants in the spring and fall, as well as for its summer resident waterfowl populations (Whitelaw 1989*),. Results (A) During the 1982-1990 period, no cormorants were observed on any of the loon study lakes after 438 visits to 94 lakes (Table 1). In contrast, 1—2 indi- viduals were observed on eight occasions from 1993 to 1998 on seven of the 94 study lakes visited 308 times (Table 1). Most sightings involved single birds on the lakes and occurred in September. The dis- tance between each of the seven lakes involved and the nearest portion of Georgian Bay varied from 7 to 160 km. The lakes closest to Georgian Bay were in — Killarney Provincial Park. (B) The first cormorant sighting reported to the SOS in the Sudbury Region was a single bird on Lake Panache on 22 May 1971 seen by Ross Lowe (Nicholson 1974*). No other cormorant sightings were reported to the SOS until 1979. Eight sightings (1-3 birds per sighting) were reported from the } Sudbury Region in 1979-1985 (Table 2). None were reported in 1986-1988 except at Kelley Lake (Figure 1 and see below). From 1989 to 1999, however, there were 17 sightings with up to 47 birds per sight- ing, excluding those reported from Kelley Lake. All these observations were made within 30 km of Sudbury (Figure 1, Table 2). Unconfirmed reports (not in the SOS files) of 2002 ALVO, BLOMME, AND WESELOH: CORMORANTS AT INLAND LAKES 361 80° James Bay 560 km Lake Abitibi @58 260 km mts Ella Islet ~~ Lake Superior 340 km _——_—————~_ °° | 0 50km _—k——_ 0 25 mi \ ee \ Lake Nipissing = Nipissing 100 km pon e Killamey Prov. f Park ‘a4, 70 km North 31 Bae B ast Channel oS ae = 46° Nene oa oe a mH Gi f acpi = ay ena ee 80° FIGURE 1. Map showing the area north of Lake Huron (Georgian Bay), including: (a) numbered sites from the “outlying Sudbury region” (delimited by dashes) (see Table 1 and Alvo (1985*)); (b) lettered sites where SOS observations were reported (see Table 2), (c): blow-up of the immediate area of the City of Sudbury; (d) location of Wanapitei Lake, (e) the direction and distance of the nearest cormorant colony in each major direction, and (f) inset showing study area in Ontario. cormorants observed on shoals and islands on the southeast corner of Wanapitei Lake since 1995 were made, and up to seven individuals were seen in flight over the lake (B. Olivier, personal communication). This is by far the largest lake in the Sudbury region (13 131 ha). In an attempt to verify these observa- tions and determine whether breeding had occurred, a low-level flight by float plane was made on 25 October 1999 over the shoals and islands. Definitive remains of at least 14 cormorant nests were observed on one of the small unnamed islands (46°43'20’N, 80°40'50”W) (DVW and L. Shutt, Canadian Wildlife Service, personal observation). (C) Neither CB nor other SOS members noted any summer resident cormorants (1.e., cormorants that appeared to have taken up extended daily residence) on Kelley Lake from 1972 to 1986, despite an annual average of 14.5 visits by CB alone (with a minimum of two visits in 1977 and a maximum of 29 in 1980). The first summer resident cormorants were reported on Kelley Lake in 1987 (Figures | and 2). From 1987 to 1999, the reported cormorant numbers on Kelley Lake fluctuated from | to 103; birds were reported there in all months from 8 April to 4 October. Breeding began there in 1996 with two nests containing five eggs (12 June) and was the first known nesting for the Sudbury 362 THE CANADIAN FIELD-NATURALIST Vol. 116 TABLE 1. Summary of (A) RA’s observations of Double-crested Cormorants on inland lakes north of Georgian Bay, 1982-1998. Date Lake No.! 1982-1990 13 September 1993 4 16 September 1993 58 19 June 1996 31 03 August 1996 34 14 September 1996 28 14 September 1997 2 19 September 1997 84 05 September 1998 + Lake Size # (ha) Cormorants No cormorants observed. @) te) N i=) i) — et LO '_Lake numbers correspond to those used by Alvo (1985*). Region (Peck 1997*). In-1997, six nests and 15 young were counted on 29 July (C. Blomme, unpublished data). The colony was active in both 1998 and 1999, when young were observed. Definitive remains of 26 nests and one dead juvenile were counted on 18 November 1999. Nesting occurred in 2000 through 2002 inclusive. Discussion With one exception, cormorants were not reported within 35-40 km of Sudbury until 1979. Birds were not “seasonal residents” in this region until 1987, and the first breeding apparently did not occur there until 1996. In the outlying Sudbury region, no cor- morants were observed during the nine-year period 1982-1990, but they were observed regularly in small numbers, primarily in the autumn, from 1993 onwards. In the late 1970s to mid-1980s, when only one to three cormorants per year were being reported in the TABLE 2. List of all observations of Double-crested Cormorants on inland lakes near Sudbury as reported by the Sudbury Ornithological Society from 1972 to 1999. Only the first sighting is listed for Kelley Lake. Observations at sites other than lakes represent birds in flight. Letters in parentheses refer to lakes mapped in Figure 1. Date Site 22 May 1971 Panache Lake (A) 1972-1978 24 June 1979 Panache Lake 06 May 1980 Bethel Lake (B) 29 April 1981 Whitewater Lake (C) 24 May 1981 Kelley Lake (D) 28 April 1982 Whitewater Lake 29 May 1982 Whitewater Lake 15 May 1985 Copper Cliff (town) (E) 15 June 1985 Sudbury (city) (F) 26 April 1989 Lively (town) (G) 27 August 1990 Ramsey Lake (H) 31 August 1990 Bethel Lake 12 May 1991 Robinson Lake (1) 29 May 1991 Meatbird Lake (J) 07 September 1991 Laurentian University 01 September 1992 Bethel Lake 01 October 1992 Bethel Lake 26 April 1995 Whitewater Lake 22 July 1995 Lively (town) 07 August 1995 Creighton Mine, Lively 17 September 1995 Halfway Lake (K) 01 September 1997 Ramsey Lake 22 April 1998 Whitewater Lake 31 May 1998 Robinson Lake 21 April 1999 Whitewater Lake 24 September 1999 Meatbird Lake Size (ha) # birds 8959 1 No cormorants observed. N id SAD DD ORK ON RR OD BOW DN FB WD BB BOD (town) — ~ \o AS 2002 Sudbury Region, the nearest known breeding colonies were in northern Georgian Bay and the North Channel of Lake Huron, 70-100 km away (Figure 1). Colonies there were growing rapidly. In northern Georgian Bay nest numbers (= pairs of breeders) increased from 64 to 342 in 1980-1985 — an annual growth rate of 40% (DVW unpublished data). Other active breeding sites in northern Ontario at that time may have included Rupert Bay in southern James Bay, more than 500 km north, where breeding occured in 1912, the 1940s (200 nests) and 1995 (42 active nests in 1995; Tymstra 1997), and Lake Abitibi, 260 km northeast, where undetermined numbers bred since at least 1982 (Chuck McCrudden, personal communication). Thus, sightings in the Sudbury Region within 35-40 km of “” EllO e@ Observations ve) o) —- NN Aw hbAaA Ww Ooonownouw Oo wn lO NUMBERS OF DOUBLE CRESTED CORMORA I986 87 88 89 90 OI ALVO, BLOMME, AND WESELOH: CORMORANTS AT INLAND LAKES earliest date Aprill7,I99I latest date October 13, I997 363 the city during the late 1970s to mid-1980s, which were exclusively in the spring, could have been of birds from Georgian Bay that had wandered inland or “overshot” their migratory destination. Alternatively, they could have been migratory birds en route to James Bay or Lake Abitibi. The lack of resident birds suggests their stays were short and that they went (returned) elsewhere to breed. Cormorants were first “resident” in the Sudbury Region at Kelley Lake in 1987-1989, when 1—2 birds remained each summer. Summer numbers increased annually after 1990 until breeding occurred in 1996 (C. Blomme, unpublished data). The increasing num- ber of non-breeders at Kelley Lake, April—October, 1990-1996, may have resulted from wandering imma- elepteeeSept2 ® ee & @ GO tone xe evunel2 @ e "s equuly2s 92 93 94 95 96 97 98 99 OO TIME OF OBSERVATIONS ON KELLEY LAKE , SUDBURY FIGURE 2. Plot of numbers of Double-crested Cormorants seen on a given date between 1986 and 1999 at Kelley Lake, Sudbury, Ontario. Maxima of over 100 birds were seen on 22 September 1994 and 2 September 1996 and represent migrants as well as resident birds. 12 June 1996 and 29 July 1997 show nest confirmation dates. 364 ture birds, adults which failed as breeders elsewhere or that, for some reason, did not breed at another colony site, or additional birds overshooting their Lake Huron destinations. It is unlikely they were actively breeding birds from other colonies as they would not have taken up residence away from their nesting colony. Kelley Lake thus became a population centre for non-breeding birds. During this time, colonies in northern Georgian Bay continued to grow, from 840 nests in 1990 to 3000 nests in 1997 (DVW, unpublished data), so there would have been ample immature birds and inexperienced adult birds looking for nest sites. Cormorants are often “resident” at a site several years before they eventually breed there (Hatch and Weseloh 1999). In the late 1980s and early 1990s, new colonies continued to appear in northern Ontario. They were discovered at: Ella Islet, Lake Superior, near Wawa, 340 km NW of Sudbury, with two nests in 1989 (Blokpoel and Tessier 1993) and 138 nests in 1999 (DVW, unpublished data), and Lake Nipissing, 70 km east of Sudbury, in 1993 (and possibly earli- er); numbers there grew from 638 to 1600 nests from 1995 to 1999 (26%/yr; Richard Rowe, personal com- munication). Immature and inexperienced adults tak- ing up residence at Kelley Lake might have come from these colonies. In the early 1990s the number of sightings in the Sudbury Region increased (Table 2). These birds were probably dispersing locally from Kelley Lake to other lakes to feed, as those lakes were within the usual daily foraging range for cormorants, up to 30 km (Custer and Bunck 1992). In 1993, cormorants first started being observed on lakes in the outlying Sudbury region, primarily in August and September (data set A). Birds at lakes close to Georgian Bay (i.e., lake numbers 31 and 34) probably came from the Bay, as the flight distances would be <5 km. Alternatively, they could have been migrants from farther north, including Kelley Lake and Wanapitei Lake. Birds observed at lakes farther from Georgian Bay (i.e., lake numbers 2, 4, 28, 58 and 84) were more likely to be from Wanapitei or Kelley Lakes. Alternatively, they could have been autumn migrants from breeding sites farther north, but given the ori- entation of the rivers to the north, they would have been autumn migrant stragglers or other wandering birds. It is unlikely these birds came from Georgian Bay, as cormorants are not known to make north- ward post-fledging dispersal as do some herons (Dolbeer 1991). Birds reported in this study (especially prior to summer residency being established at Kelley Lake) were not observed to fly inland but rather “appeared” or were found at inland sites. In almost all cases, these inland sites were far enough removed from the “large lakes” that the cormorants’ appearance at them was probably not the result of foraging flights from a THE CANADIAN FIELD-NATURALIST Vol. 116 breeding colony. Thus there are at least two kinds of inland occurrences of cormorants. The first involves stragglers that either strayed off a migratory route or were not attached to a colony and were wandering. If, in any given area, the number of these individuals gradually increases, new colonies may form. This appears to fit most of the birds seen in early years of this study and may be how the colonies at Kelley and Wanapitei Lakes were formed. The second type of inland occurrence is that of birds dispersing inland on a daily basis from a breed- ing colony or some other roosting situation. This probably accounts for many of the sightings in the immediate Sudbury area once cormorants had estab- lished summer residency at Kelley Lake; i.e., birds left Kelley Lake each morning to feed in the sur- rounding area, and returned to Kelley to roost. This is also a type of behaviour that has been reported at sev- eral Great Lakes coastal cities/towns where cor- morants have been observed flying inland along river courses; e.g., in Ontario at Kingston (DVW, personal observation), Parry Sound (R. Black, Ontario Ministry of Natural Resources), Blind River (S. Elliott, Ontario Ministry of Natural Resources) and in New York near Watertown (I. Mazzocchi, New York State Department of Environmental Conservation). These flights may or may not lead to formation of new colonies depending upon the status of the birds involved. Birds with a firm position at a breeding colony (i.e., a mate, nest and eggs or young), would have a strong motivation to return to their home colony. Non-breeding birds or perhaps newly estab- lished birds at the periphery of the colony could be attracted to new inland sites by the prospect of reduced intra-specific competition. What impact do cormorants have on inland wildlife? Long-term monitoring of presence and breeding on inland waters is needed in the Sudbury region. Inland waters in other portions of the Great Lakes watershed should be searched. Research into the impacts on other birds and on fish in the Sudbury region could have implications throughout the Great Lakes watershed. Radio-tracking could be used to determine the origin of the birds using small inland lakes. Acknowledgments This project was supported by the Helen McCrae Peacock Foundation, the Canadian Wildlife Service (Ontario Region), World Wildlife Fund (Canada), the Canadian Wildlife Federation, the Ontario Ministry of Natural Resources, the North American Loon Fund, and the James L. Baillie Memorial Fund. We thank the many contributors to the Sudbury Ornithological Society’s monthly reports, especially Gloria Blomme, Chris Bell, Charlie Whitelaw, Heather Baines, Don Ferguson, and the editors John Nicholson and John Lemon. We thank Richard Rowe, David Fluri, and Chuck McCrudden of the 2002 Ontario Ministry of Natural Resources, and Nathalie Tessier, George Lyons and Brenda Harrow for pro- viding valuable information. Phil McColl of the Drafting Department at the Canada Centre for Inland Waters, Burlington, Ontario, prepared the map. A. J. Erskine, Josée Nesdoly; Cynthia Pekarik; and an anonymous reader reviewed the manuscript. Documents Cited [marked * in text citation] Alvo, R. 1985. The breeding success of Common Loons (Gavia immer) in relation to lake alkalinity. M.Sc. the- sis, Trent University, Peterborough, Ontario. 122 pages. Nicholson, J. C. 1974. The birds of the Sudbury District: a species accounts summary. Sudbury, Ontario. 57 pages. Peck, G. K. 1997. Ontario nest records scheme, twenty eighth report (1956-1996). Toronto, Ontario. 21 pages. Postupalsky, S. 1978. Toxic chemicals and cormorant populations in the Great Lakes. Canadian Wildlife Ser- vice. Wildlife Toxicology Division. Manuscript Report Number 40, Ottawa. Whitelaw, C. 1989. Seasonal occurrence of birds in the Sudbury District. Sudbury Ornithological Society. Sudbury, Ontario. 10 pages. Wires, L., and F. Cuthbert. 1999. Status of the Double- crested Cormorant (Phalacrocorax auritus) in eastern and central North America. Draft report to the U.S. Fish and Wildlife Service. Fort Snelling, Minnesota. 152 pages. Literature Cited Alvo, R. 1996. Double-crested Cormorant. Pages 232-235 in The breeding birds of Québec: Atlas of the Breeding Birds of Southern Québec. Edited by J. Gauthier and Y. Aubry. Association québécoise des groupes d’ornitho- logues, Province of Québec Society for the Protection of Birds, Canadian Wildlife Service, Environment Canada, Québec Region, Montréal. 1302 pages. Alvo, R., D. J. T. Hussell, and M. Berrill. 1988. The breeding success of common loons (Gavia immer) in relation to alkalinity and other lake characteristics in Ontario. Canadian Journal of Zoology 66: 746-752. Blokpoel, H., and G. D. Tessier. 1993. Atlas of colonial waterbirds nesting on the Canadian Great Lakes, 1989-1991. Part 1. Cormorants, gulls and island-nesting terns on Lake Superior in 1989. Technical Report Series (181). Canadian Wildlife Service, Ontario Region. 96 pages. Blokpoel, H., and G. D. Tessier. 1997. Atlas of colonial waterbirds nesting on the Canadian Great Lakes, 1989-1991. Part 2. Cormorants, gulls and island-nesting terns on Lake Huron in 1989. Technical Report Series (259), Canadian Wildlife Service, Ontario Region. 154 pages. Christie, W. J. 1974. Changes in the fish species compo- sition of the Great Lakes. Journal of the Fisheries Research Board of Canada 31: 827-854. Christie, W. J., K. A. Scott, P. G. Sly, and R. H. Strus. 1987. Recent changes in the aquatic food web of eastern ALVO, BLOMME, AND WESELOH: CORMORANTS AT INLAND LAKES 365 Lake Ontario. Canadian Journal of Fisheries and Aquatic Sciences 44 (Supplement 2): 37-52. Custer, T. W., and C. Bunck. 1992. Feeding flights of breeding Double-crested Cormorants at two Wisconsin colonies. Journal of Field Ornithology 63: 203-211. Dolbeer, R. A. 1991. Migration patterns of Double-crest- ed Cormorants east of the Rocky Mountains. Journal of Field Ornithology 62: 83-93. Hatch, J. J. 1984. Rapid increase of Double-crested Cormorants nesting in southern New England. American Birds 38: 984-988. Hatch, J. J., and D. V. Weseloh. 1999. Double-crested Cormorant (Phalacrocorax auritus). In The Birds of North America (441) Edited by A. Poole and F. Gill. The Birds of North America, Inc., Philadelphia, Pennsyl- vania. Ryckman, D. P., D. V. Weseloh, P. Hamr, G. A. Fox, B. Collins, P. J. Ewins, and R. J. Norstrom. 1998. Spatial and temporal trends in organochlorine contami- nation and bill deformities in Double-crested Cormor- ants (Phalacrocorax auritus) from the Canadian Great Lakes. Environmental Monitoring and Assessment 53: 169-195. Scharf, W.C., and G. W. Shugart. 1981. Recent increas- es in Double-crested Cormorants in the United States Great Lakes. American Birds 35: 910-911. Schneider, C. P., A. Schiavone Jr., T. H. Eckert, R. D. McCullough, B. F. Lantry, D. W. Einhouse, J. R. Chrisman, C. M. Adams, J. H. Johnson, and R. M. Ross. 1998. Double-crested Cormorant predation on smallmouth bass and other warm water fishes of the eastern basin of Lake Ontario: Overview and summary. New York State Department of Environmental Conser- vation Special Report. 6 pages. Taverner, P. A. 1919. Birds of eastern Canada. Geolog- ical Survey, Department of Mines, Memoir 104, Number 3, Biological Series. Ottawa. 297 pages. Tymstra, Y. R. 1997. Bird observations from some south- eastern James Bay Islands, N.W.T. Ontario Birds 15: 108-115. Weseloh, D. V., S. Brechtel, and R. D. Burns. 1979. Recent population changes in Double-crested Cor- morants and California and Ring-billed Gulls in Alberta, Canada, with a note on White Pelicans. Pages 10-18 in Proceedings of the 1977 Conference on Colonial Waterbird Group. Weseloh, D. V., and B. Collier. 1995. Great Lakes Fact Sheet: The rise of the Double-crested Cormorant on the Great Lakes: winning the war against contaminants. Minister of the Environment and Public Works and Government Services Canada. 12 pages. Weseloh, D. V., P. J. Ewins, J. Struger, P. Mineau, C. A. Bishop, S. Postupalsky, and J. P. Ludwig. 1995. Double-crested Cormorants of the Great Lakes: changes in population size, breeding distribution and reproduc- tive output between 1913 and 1991. Colonial Waterbirds 18 (Special Publication 1): 48-59. Received 27 April 2000 Accepted 13 June 2002 Seasonal Distribution of Waterbirds in Relation to Spawning Pacific Herring, Clupea pallasi, in the Strait of Georgia, British Columbia TERRANCE M. SULLIVAN!, ROBERT W. BUTLER2, and W. SEAN Boyb2 13977 Nicomek] Road, Surrey, British Columbia, V4P 2J9 Canada 2Canadian Wildlife Service, Pacific Wildlife Research Centre, RR1, 5421 Robertson Road, Delta, British Columbia V4K 3N2 Canada Sullivan, Terrance M., Robert W. Butler, and W. Sean Boyd. 2002. Seasonal distribution of waterbirds in relation to Spawning Pacific Herring, Clupea pallasi, in the Strait of Georgia, British Columbia. Canadian Field-Naturalist 116(3): 366-370. About 3500-3700 waterbirds were present in 23 bays and beaches along 150 km of shoreline on the east coast of Vancouver Island, British Columbia prior to the arrival of Pacific Herring (Clupea pallasi). Waterbird abundance in the region increased to > 32 500 individuals when herring spawned in March with the greatest proportional increase occurring at the spawn site. Another 140 000 waterbirds were present outside the study site in deep water a few km offshore. Waterbird density in the entire study area increased from a low of about 66 birds/km2 prior to the arrival of herring to a peak density of about 616 birds/km? when the herring spawned on the beaches. Key Words: Loons, grebes, seabirds, sea ducks, gulls, Pacific Herring, Clupea pallasi, Strait of Georgia, British Columbia. From February to April, large flocks of birds are attracted to schools of Pacific Herring (Clupea pallasi Valenciennes 1847) that spawn in shallow waters along the Pacific coast of Canada (Munro and Clemens 1937; Haegele and Schweigert 1989; Haegle 1993; Vermeer et al. 1997). One of the largest spawn- ing sites in western Canada is on the western shore of the Strait of Georgia in southern British Columbia. Herring enter the Strait of Georgia in early winter to deposit eggs in late February and early March (Hay et al. 1989). The eggs hatch about 14 days after they are deposited and the larvae form into schools that depart for deep water in April (Haegele and Schweigert 1989). At least 21 waterbird species eat Pacific Her- ring or their eggs in British Columbia including Western Grebe (Aechmophorus occidentalis), Brandt’s Cormorant (Phalacrocorax pencillatus) and Pelagic Cormorant (Phalacrocorax pelagicus), Brant (Branta bernicla), Black Scoter (Melanitta nigra), Surf Scoter (Melanitta perspicillata), White-winged Scoter (Melannita fusca), Harlequin Duck (Histrionicus histrionicus), Long-tailed Duck (Clangula hyemalis), Greater Scaup (Aythya marila), Common Goldeneye (Bucephala clangula), Barrow’ s Goldeneye (Bucephala islandica), Bufflehead (Bucephala albeola), Common Merganser (Mergus merganser), Red-breasted Merganser (Mergus serra- tor), Surfbird (Aphriza virgata), Black Turnstone (Arenaria melanocephala), Glaucous-winged Gull (Larus glaucescens), Mew Gull (Larus brachyrhyn- chos), California Gull (Larus californicus), and Marbled Murrelet (Brachyramphus marmoratus) [Grosz and Yocom 1972; Haegele 1993; Vermeer et al. 1997; RWB, personal observation]. Large numbers of waterbirds attracted to beaches with spawning Pacific Herring in British Columbia (Vermeer 1981; Haegele 1993; V Grmceniaaaer 1997) are potentially at risk to a single oil spill and to commercial exploitation of herring. The numbers of waterbirds attracted to spawning sites originate from a wide but undocumented area. It is also unknown how different species utilize the spawn. Here, we describe the magnitude of regional shifts in abundance of waterbirds at one spawn site in relation to the arrival of adult Pacific Herring, the presence of herring eggs and the emergence of lar- val herring. Methods and Study Site The Strait of Georgia is a 200 km long body of water separating Vancouver Island from the mainland of British Columbia (Vermeer and Butler 1989). The east coast of Vancouver Island includes many small bays with rocky, gravel or sand substrates frequented by sea ducks, loons, grebes, shorebirds, and seabirds in fall, winter and spring. Each spring, Pacific Herring spawn in Lambert Channel and neighboring bays in the Strait of Georgia (Figure 1). We counted waterbirds in 23 bays and beaches between Nanaimo and Campbell River including Lambert Channel, once a week for 10 weeks between 19 February and 23 April 1998 (Figure 1). The sites were Nanaimo estuary, Departure Bay, Piper’s Lagoon, Oar Road, Snaw-na-as dock, Nanoose Bay, Rathrevor Beach, Parkville Beach, Qualicum Beach, Qualicum Bay, Deep Bay, Fanny Bay, Buckley Bay, Fillongley Park, Hornby Ferry Terminal, 1565, Union Bay, Royston Beach, Comox Harbour, Comox Ferry Terminal, Oyster River, Willow Point, and Campbell River. All surveys were made from land through 8 power binoc- 366 2002 SULLIVAN, BUTLER, AND BOYD: DISTRIBUTION OF WATERBIRDS 367 Campbell River e — 2 = Cal s Baynes ©, Sound “~~, eo. 0 2s _Lambert e@ Channel Vancouver S&S o.. GX FIGURE |. Surveyed area on Vancouver Island showing the location of 23 beaches included in this survey and the major Pacific Herring spawning site in Lambert Channel in 1998. ulars and a 15-45 X or 20-60 telescope. All birds seen on or flying over the water within approximately 1 km of shore were tallied to species on most days and grouped by genus or family when identification to species was difficult as a result of poor visibility on windy days. Herring were visible by the presence of ripples on the water surface when schools moved close to shore, and spawning was evident when the water turned a milky color from the release of sperm. Herring eggs require about 14 days to hatch (Haegele and Schweigert 1989). We divided our waterbird censuses into five time periods to coincide with stages of the herring spawning season as follows: (1) the pre- spawn stage was defined as the period before herring schools were first seen in the study area (< 26 February), (2) the adult fish stage was estimated from when herring were first observed until the day before spawning began (27 February to 12 March), (3) the spawning stage occurred from when sperm was first noted until 14 days later when the first eggs were esti- mated to have hatched (13 to 27 March), (4) the early larval stage was estimated to begin two weeks after the start of the spawning period and ran for a further two weeks (i.e., >27 March—9 April), and (5) the late larval stage ran from 10-23 April. We estimated the surveyed area of each of the 23 sites (Figure 1) by the proportional area of water within 1 km radius of the survey point. Results A total of 93 703 birds representing 28 species or families of birds were counted over all surveys (Table 1). The four most abundant taxa were scoters (36.3%), gulls (18.2%), Brant (12.4%) and scaup (11.7%). The number of waterbirds present in the study area during the herring spawn seasons exceed- ed the late winter pre-spawn counts for at least 49 days. During the pre-spawn stage, we estimated between 3507 and 3718 waterbirds, of which over half were scoters (55.5%), frequented the bays and beaches in the study area (Table 2). Nearly 21 000 birds were present during the adult fish stage and over 32 500 birds were present at the end of the spawning period (Table 2). The number of water- birds declined to between 6 991 and 13 325 individ- uals during the herring larval stages (Table 2). The influx in waterbirds was noticeable through- out the study area but the greatest proportional increase occurred at the major spawn site in Lambert Channel. This channel held only about 200 birds prior to the arrival of herring but abundance grew to 368 TABLE |. Total number of waterbirds counted during 10 surveys of the east coast of Vancouver Island between Nanaimo THE CANADIAN FIELD-NATURALIST and Campbell River, British Columbia, 19 February to 23 April 1998. Vol. 116 Species Scientific Name Number % Common Loon Gavia immer 894 1.0 Pacific Loon Gavia pacifica 1776 LS Western Grebe Aechmophorus occidentalis 765 0.8 Red-necked Grebe Podiceps grisgena 1024 1.0 Horned Grebe Podiceps auritus 803 0.9 Double-crested Cormorant Phalacrocorax auritus 219 0.2 Pelagic Cormorant Phalacrocorax pelagicus 187 0.2 Trumpeter Swan Cygnus buccinator 8 <0.1 Brant Branta bernicla 11648 12.4 Canada Goose Branta canadensis 4] <0.1 Mallard Anas platyrhynchos 731 0.8 American Wigeon Anas americana 3728 4.0 Northern Pintail Anas acuta 340 0.4 Dabbling Duck spp.? Anas spp. 4081 4.4 Greater Scaup Aythya marila 1701 1.8 Lesser Scaup Aythya affinis 31 <0.1 Scaup spp.> Aythya spp. 9269 aS White-winged Scoter Melanitta fusca 3400 3.6 Black Scoter Melanitta nigra 2187 yi Surf Scoter Melanitta perpicillata 11146 Lin Scoter spp.° Melanita spp. 17322 18.5 Harlequin Duck Histrionicus histrionicus 613 0.7 Long-tailed Duck Clangula hyemalis 687 0.7 Barrows Goldeneye Bucephala islandica 50 <0.1 Common Goldeneye Bucephala clangula 801 0.9 Bufflehead Bucephala albeola 2032 pigs Common Merganser Mergus merganser 42 <0.1 Red-breasted Merganser Mergus serrator 901 1.0 Gull spp.4 Larus spp. 17021 18.2 Pigeon Guillemot Cepphus columba 32 <0.1 Marbled Murrelet Brachyramphus marmoratus poe 0.2 Total 93 703 aAnas platyrhynchos and A. americana. most likely Aythya marila. ‘Melanitta nigra, M. perspicllata, and M. fusca. dZarus glaucescens, L. californicus, L. brachyrhynchos. over 12 600 birds by the end of the spawn stage (Table 2). Many more birds were present than our surveys indicated. One estimate of the offshore flocks made from a boat in late March reported about 140 000 waterbirds (I. Goudie, CWS, Delta, personal communication). The seasonal significance of herring spawn to the presence of waterbirds on the east coast of Vancouver Island is illustrated well by the censuses in Lambert Channel. The density of birds was about 80 times greater when herring were present than prior to their arrival (Table 3). Soon after the herring spawned, the number of waterbirds plummeted in Lambert Channel but they were still more abundant than during the pre-spawn period. The predominant waterbirds prior to the arrival of herring in Lambert Channel were scoters (82.8%) and the Harlequin Duck (14.5%). Gulls (76.7%) fol- lowed by scoters (16.3%) were the most numerous birds during the adult fish stage (Table 3). We did not count the gulls in the pre-spawn stage although we noted small numbers. Scoters (43.5%), gulls (20.6%) and Long-tailed Duck, cormorants and scaup (7.9-9.7%) predominated during the spawn stage. The herring early larval stage was predominat- ed by a large influx of migrating Brant (50.6%), gulls (26.3%), and Bufflehead (12.0%). Scoters (42.6%), Brant (29.9%) and Pacific Loons (19.6%) were the predominant avifauna during the late larval stage. The occurrence of many species of waterbirds in Lambert Channel coincided with the herring repro- ductive stage that matched their particular food pref- erence. The Common Loon was significantly (P< 0.001) more numerous when adult Herring were present. Cormorants, scaup, scoters, Common Goldeneye, Long-tailed Duck, and gulls were signifi- cantly (P< 0.001) more numerous in late March when Herring were spawning (Table 3). The number of grebes present during the spawn and early larval stages was not significantly different (P > 0.05) but they were significantly (P< 0.01) more numerous during the early larval stage than the late larval stage. 2002 SULLIVAN, BUTLER, AND BOYD: DISTRIBUTION OF WATERBIRDS 369 TABLE 2. Total number and density (birds/km2) of waterbirds counted at 20 sites on the east coast of Vancouver Island and three sites in Lambert Channel, 19 February—23 April 1998. Number (density) of birds Number (density) Spawning Vancouver Island of birds at Lambert Number (density) stage? Date sites Channel at all sites Pre-spawn 19 February 3492 (68.1) 226 (144.0) 3718 (70.4) 24 February 3 313.' (64.6) 194 (123.6) 3507 (66.4) Adult fish 05-06 March 9295 (181.2) 4218 (2686.6) 13°513 42559) 11-12 March 4953 (96.6) 16 012 (10198.7) 20965 (397.1) Spawn 18-19 March 1515930.. (302.7) 10487 (6679.6) 26018 (492.8) 25-26 March 19 860 (387.1) 12661 (8064.3) 32 521 (615.9) Early larval 01-02 April 10150 (197.9) 1256 (800.0) 11 406 (216.0) 08-09 April 6 253° (121.9) 738 (471.3) 6991 (132.4) Late larval 16-17 April 8 684 (169.3) 953 (607.0) 9537 (180.6) 23 April 12992 (237.3) 1153 (734.4) 13 325 (252.4) Total 93 703 (1826.6) 47 899 (30509.6) 141 601 (2681.9) apre-spawn stage < 26 February; adult fish stage 27 February to 12 March; spawning stage 13 to 27 March; early larval stage 27 March—9 April; and late larval stage 10—23 April The number of Brant was significantly smaller during the spawning stage than either the early or late larval stages. Discussion The arrival of herring along the coast of British Columbia in late winter and spring creates a seasonal bonanza of food for thousands of waterbirds (Munro and Clemens 1937; Haegele and Schweigert 1989; Haegele 1993; Vermeer et al. 1997). The high densi- ty of birds reported indicated that many species moved to herring spawn sites but the distances they traveled was unknown. Our data showed that the seasonal increase in abundance was many times greater than the pre-spawn estimates in the study area indicating that many waterbirds might have traveled more than 80 km to eat herring in Lambert Channel. It is unlikely that the large flocks counted in Lambert Channel were the cumulative total from bays we did not survey between Nanaimo and Campbell River because we recorded increased num- bers of waterbirds in all bays, including Lambert Channel. The number of waterbirds present in the study area was greater when herring spawned than during the pre-spawn period and many more settled offshore. The combined single offshore estimate of 140 000 waterbirds a few kilometers outside our study area and our peak shoreline estimate of 32 521 waterbirds in the study area in late March (Table 2) suggest that about 170 000 waterbirds might have been present during the Herring spawn event in 1998. Compared with the 3507 waterbirds present TABLE 3. Number of waterbirds in Lambert Channel prior to the arrival of Pacific Herring, and during the adult fish, egg, and early and late larval Pacific Herring stages. nd = no data. Different superscripts indicate significant differences between stages (P< (0.05). Adult Species Pre-spawn fish Common Loon 0 1684 Pacific Loon 1 0 Red-necked Grebe 0 0 Horned Grebe 2 0 Cormorant spp. 3 1504 Brant 0 1464 Scaup spp. 0 9154 Scoter spp. 188 3 2684 Harlequin Duck 33 ri Long-tailed Duck 0 0 Common Goldeneye 0 0 Bufflehead 0 0 Red-breasted Merganser 0 0 Gulls nd 15 3504 Total (all birds) 227 19 997 Spawn Early larval Late larval 248 6c 22 12 0 406 5A 64 0 17A 234 8B 2 0008 4c 8 4468 974¢ 618P 1 8358 &C 0 10 0598 33¢ 881 0 0 0 22344 7B 0 9684 418 0 6614 2308 13¢ 97 85 72 47758 506¢ 39D 23 133 1923 2 067 370 prior to the spawn, this indicates a 48-fold increase in waterbirds on the east coast of Vancouver Island. Our observations generally supported our hypoth- esis that the timing of arrival of waterbirds coincid- ed with the presence of their diet preferences. Loons were most numerous when adult fish were schooling offshore in deepwater, gulls, grebes and cormorants pursued adult fish when they entered the shallows to spawn and diving ducks gorged on herring eggs. Vermeer et al. (1997) and Reed et al. (1998) sug- gested that Brant intentionally ingest herring eggs clinging to Eelgrass (Zostera marina) and Haegele (1993) reported Brant eating herring eggs from the beaches in Lambert Channel, which we also observed in our study. Brant occur on beaches with herring spawn in significantly larger flocks than in areas without spawn (Vermeer et al. 1997). How- ever, the number of Brant was significantly smaller during the spawning stage when eggs were most numerous on eelgrass than either the early or late larval stages. Waterbirds are reported to consume a very small proportion of the total egg mass laid by herring (Haegele and Schweigert 1989; Haegele 1993) but the importance of herring as a source of food energy for waterbirds has not been quantified. Our study was not designed to examine this relationship specif- ically. Loons, grebes, and waterfowl that have been reported to occur at herring spawn events, migrate from the Strait of Georgia during or soon after her- ring depart for deep water in late March and early April (Campbell et al. 1990). Herring, eggs, and lar- vae could possibly provide for a rapid gain in lipids that waterbirds might use to power migratory flights to other herring spawning beaches or to the breeding grounds. The migrations of several shorebirds coin- cide with a seasonal abundance of herring eggs and invertebrate prey (Gratto et al. 1984; Hicklin 1987; Norton et al. 1990; Clark et al. 1993). If our hypoth- esis is correct, then the fitness of many waterbirds along the east coast of Vancouver Island and perhaps other regions of the Strait of Georgia and Puget Sound may be closely linked to the quantity and duration eggs and Herring are available to them on beaches in spring. Acknowledgments We thank Ian Goudie for providing us with addi- tional census data. This study was funded in part by the Pacific Coast Joint Venture under the auspices of the Canadian Wildlife Service. Fred Cooke, John Elliott and Anthony Erskine commented on the manuscript, and Pam Whitehead prepared the figure. THE CANADIAN FIELD-NATURALIST Vol. 116 Literature Cited Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. H. Cooper, G. W. Kaiser and M. C. E. McNall. 1990. The birds of British Columbia. Volume 1. Royal British Columbia Museum, Victoria, British Columbia. Clark, K. E., L. J. Niles, and J. Burger. 1993. Abun- dance and distribution of migrant shorebirds in Delaware Bay. Condor 95: 694 —705. Gratto, G. W., M. L. H. Thomas, and C. L. Gratto. 1984. Some aspects of the foraging ecology of migrant juvenile sandpipers in the outer Bay of Fundy. Canadian Journal of Zoology 62: 1889-1892. Grosz, T. and C. F. Yocom. 1972. Food habits of the White-winged Scoter in northwestern California. Journal of Wildlife Management 36: 1279-1282. Haegele, C. W. 1993. Seabird predation of Pacific Herring, Clupea pallasi, spawn in British Columbia. Canadian Field-Naturalist 107: 73-82. Haegele, C. W., and J. E. Schweigert. 1989. Egg loss from Pacific Herring spawns in Barkley Sound in 1988. Canadian Manuscript Report of Fisheries and Aquatic Science Number 2037. Hay, D. E., M. C. Healey, L. J. Richards, and J. B. Marliave. 1989. Distribution, abundance and habitat of prey fishes in the Strait of Georgia. Pages 37-49 in The ecology and status of marine and shoreline birds in the Strait of Georgia, British Columbia. Edited by K. Vermeer and R. W. Butler. Special Publication, Canadian Wildlife Service, Ottawa, Ontario. Hicklin, P. W. 1987. The migration of shorebirds in the Bay of Fundy. Wilson Bulletin 99: 540-570. Munro, J. A., and W. A. Clemens. 1937. Waterfowl in relation to the spawning of herring in British Columbia. Biological Board of Canada 17: 1-46. Norton, D. W., S. E. Senner, R. E. Gill, Jr., P. D. Martin, J. M. Wright, and A. K. Fukuyama. 1990. Shorebirds and Herring roe in Prince William Sound, Alaska. American Birds 44: 367-371. Reed, A., D. H. Ward, D. V. Derksen, and J. S. Seeding. 1998. Brant. The birds of North America, Number 337. Edited by A. Poole, P. Stettenheim, and F. Gill. The Academy of Natural Sciences, Philadelphia and American Ornithologists’ Union, Washington, D.C. Vermeer, K. 1981. Food and populations of Surf Scoters in British Columbia. Wildfowl 32: 107-116. Vermeer, K., and R.W. Butler. 1989. The ecology and status of marine and shoreline birds in the Strait of Georgia, British Columbia. Special Publication, Cana- dian Wildlife Service, Ottawa, Ontario. Vermeer, K., M. Bentley, K.H. Morgan, and G.E.J. Smith. 1997. Association of feeding flocks of Brant and sea ducks with Herring spawn at Skidegate Inlet. Pages 102-109 in The ecology, status and conservation of marine and shoreline birds of the Queen Charlotte Islands. Edited by K. Vermeer and K. H. Morgan. Can- adian Wildlife Service Occasional Paper Number 93, Ottawa, Ontario. Received 3 August 2000 Accepted 17 June 2002 North Atlantic Sperm Whale, Physeter macrocephalus, Strandings on the Coastlines of the British Isles and Eastern Canada JOHN C. GooLp!, HAL WHITEHEAD?, and ROBERT J. REID? Institute of Environmental Science, University of Wales Bangor, Robinson Building, Deiniol Road, Bangor, Gwynedd, LL57 2UW United Kingdom. E-mail: j.c.goold@bangor.ac.uk "Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Canada. 3SAC Veterinary Science Division, Drummondhill, Statherrick Road, Inverness, [V2 4JZ United Kingdom. Goold, John C., Hal Whitehead, and Robert J. Reid. 2002. North Atlantic Sperm Whale, Physeter macrocephalus, strand- ings on the coastlines of the British Isles and Eastern Canada. Canadian Field-Naturalist 116(3): 371-388. Sperm Whale stranding records for the British Isles and Eastern Canada were analysed to investigate short and long term stranding trends on either side of the North Atlantic Ocean. Annual stranding events across a ten year period from 1988 to 1997 showed no significant trend with time in either the British Isles or eastern Canada, although stranding events were nearly three times as numerous on the Scottish coast during this period than in any other region. Strandings during this period occurred throughout the year in Scotland, Ireland and eastern Canada, but peaked in the autumn and winter periods. Strandings on the English coast were the least numerous and occurred only in the autumn and winter. Decadal analysis of stranding events for the 20 Century as a whole showed significant exponential increases in Sperm Whale strandings in all regions of the British Isles (except England due to small sample size). Decadal stranding events were fitted to an exponen- tial model which revealed a rate of increase of 14%/yr for the British Isles as a whole, beginning at about 1970. The strand- ing rate on the Scottish coast, 18%/yr, was twice that for Ireland, 9%/yr, with most of the strandings increase occurring in the offshore Scottish Islands. The time series data for eastern Canada was of insufficient extent to conduct a rigorous decadal analysis. Almost all stranded Sperm Whales on the British and East Canadian coasts have been males. Sperm Whale strandings have been most dense, and have increased fastest, in the offshore Scottish islands of Hebrides, Orkney and Shetland. The data support no firm conclusions but valid hypotheses include increased reporting and anthropogenic effects, which may be acting synergistically. The increase in the British data is too dramatic to have been caused solely by a simple increase in Sperm Whale population size. Key Words: Sperm Whale, Physeter macrocephalus, stranding, Canada, British Isles, noise, pollution, seismic, Atlantic. A number of widely publicised Sperm Whale (Physeter macrocephalus Linnaeus 1758) strandings along the coastline of the British Isles (see Geo- graphic Terminology below) in recent years has raised the profile of these events and given rise to much speculative comment. There is a general per- ception that strandings of Sperm Whales are increas- ing although the evidence is seldom critically exam- ined. Previous works indicate an increase in the number of reported strandings on the British coast- line (Berrow et al. 1993; Evans 1997), although the causes of such a trend remains elusive. If Sperm Whale strandings are on the increase, then why? Possible causes include: increased reporting of a steady stranding rate; increased density of Sperm Whales in the area of the British Isles; and increased mortality of Sperm Whales per capita of population. Such possible causes are not mutually exclusive, and could be operating synergistically if, for example, changes in migration patterns have increased the exposure of animals to anthropogenic threats. In this paper we examine geographic and tempo- ral patterns of Sperm Whale strandings in both British and eastern Canadian waters in order to quantify stranding trends. Different causes of an increase would imply different geographic and tem- poral patterns of strandings, and therefore we are able to make some progress in narrowing down potential causes. For instance, as all Sperm Whales in the North Atlantic are thought to belong to the same population (Donovan 1991), a general increase in population should be reflected on both sides of the Atlantic and be consistent with models of Sperm Whale population dynamics. Our analysis initially considers the most recent strandings on the British and East Canadian coastlines. We then relate these recent strandings to historical records as documented in the literature, and consider poten- tial factors relevant to stranding trends. Further, given the disparate archiving of data that currently exists, which is often difficult to access, we present the raw strandings data in appendices as a single, easily referenced repository of knowledge for future generations. Geographic Terminology British Isles: The land masses comprising England, Scotland, Wales, Northern Ireland and the Republic of Ireland. Atlantic Frontier: That part of the Atlantic Ocean north and west of the British Iles. Eastern Canada: The provinces of Newfoundland & Labrador, Nova Scotia, New Brunswick, Prince Edward Island, and Quebec. 371 372 Methods Regional Collection of Stranding Data Since 1913 a strandings reporting scheme has been operated by the British Natural History Museum in London. Strandings of all species have been recorded since that time, although reporting has been oppor- tunistic. In recent years, responsibility for the collec- tion of cetacean stranding data in the British Isles has been more formalised under the auspices of four regional bodies. In England, the London Natural History Museum continues to archive cetacean stranding records and, through an affiliation with the Institute of Zoology, has the remit of post-mortem examination of stranded carcasses. In the principality of Wales, a designated strandings co-ordinator affili- ates with veterinary pathologists to obtain compara- ble information; with strandings and post-mortem data from the principality ultimately archived along with the English records at the Natural History Museum (NHM). In Scotland, these tasks are under- taken by the SAC Veterinary Science Division, from where one of the authors (Reid) conducts post mortem examination of stranded cetaceans. The England and Wales stranding scheme was put on a statutory footing in 1990 with funding from the Department of the Environment (now DETR). The Scottish scheme attained similar status in 1992. Cetacean strandings data in Ireland are compiled by the Irish Whale & Dolphin Group (IWDG), with post mortem examination of beached carcasses carried out by the University College, Cork. Cetacean stranding data in Eastern Canada are collected by volunteer stranding groups in each of the five provinces bordering the Atlantic (Newfoundland and Labrador, Nova Scotia, New Brunswick, Prince Edward Island and Quebec). These are usually linked to provincial museums, local universities and/or the federal Department of Fisheries and Oceans. In the special case of Sable Island, Nova Scotia, which has one of the highest rates of cetacean strandings in the world (Lucas and Hooker 2000), a resident naturalist (Z. Lucas) keeps records of all events. Very low human population densities probably mean that there is substantial undereporting of strandings in the northern part of the Sperm Whales’ range off eastern Canada, espe- cially Labrador. Analysis of Strandings Data Recent Data Recent data for Britain and eastern Canada (1988 to 1997 inclusive) have been collated in this exercise and treated by region as numerical records showing numbers of animals stranded per year and number of stranding events per year (i.e., two or more whales stranding together are considered as one event). These events are plotted on charts to give an indica- tion of their geographic distribution around the respective coastlines. Correlation analysis of strand- THE CANADIAN FIELD-NATURALIST Vol. 116 ing events vs. year is applied to determine whether temporal trends (i.e., increase or decrease with time) exist within the initial ten year period under investi- gation for any given region. The raw data are further tabulated to show seasonality of strandings with number of stranded animals and the associated events grouped into quarterly periods. Historical Data Historical strandings data, from the official publi- cations of NHM archive data and IWDG data, are then considered and used to set the more recent data within an extended temporal context. British strand- ing records are available from 1913 in the case of England, Wales and Scotland (Harmer 1927; Fraser 1934, 1946, 1953, 1974; Sheldrick 1989; Sheldrick et al. 1994), and 1901 in the case of Ireland (Berrow and Rogan, 1997). There are no extensive historical records from eastern Canada, although records are available for Sable Island, Nova Scotia, from 1970 onwards (Lucas and Hooker 2000; Lucas, personal communication). However, records prior to 1987 were collected inconsistently and hence a decadal analysis is not attempted here, even for Sable Island. For historical comparison, the strandings data, (i.e., numbers of animals and events) are summed within ten year periods working retrospectively from the date of the most recent available data; this defines the most recent decade as the period 1988- 1997 inclusive. The data are tabulated in this decadal format and regional trends plotted. Statistical mod- elling of the trends is performed using non-linear regression, and is only performed on the events data, as this analysis is not valid for numbers of animals per se due to non independence. Only decadal peri- ods for which observations are complete are anal- ysed (1.e., the 1908-1917 decade for Scotland and England is excluded — for consistency this decade is also excluded for Ireland). Finally, the decadal stranding data for the remote Scottish Islands are separated from that of mainland Britain to yield insight into whether remote locations and the associated human communications may have led to a differential rate of reporting over historic time, and/or whether anthropogenic factors might differentially affect the outer islands. Statistical modelling Rates of increase in stranding events were calcu- lated by fitting (using least-squares) the following exponential model to the number of stranding events, n', in each year, i (using non-linear regression in SYSTAT): n,=k(1 + b%~))+error (equation 1) This models an exponential increase from a basal rate of stranding events per year k, taking off at approximately year c, and then increasing at a rate b per year. 2002 GOOLD, WHITEHEAD, AND REID: SPERM WHALE STRANDINGS 373 | | lreland Canada Sable Island (60) FiGuRE |. Distribution of Sperm Whale stranding events around the British and Eastern Canadian coastlines, 1988-1997 inclusive. Scotland (A); England & Wales (B); Ireland (C); and Eastern Canada (D). 374 Results Geographic Distribution Figure 1 shows the geographic distribution of the most recent decade (1988 to 1997) of Sperm Whale strandings, around the British and eastern Canadian coastlines. In the case of Scotland [Figure 1A], where strandings are most numerous, there is a clear tendency for strandings to occur on the western mar- gins of the Scottish mainland as well as the Outer Hebrides, Orkney Islands and Shetland Islands. There are some clear instances, from Appendix A and Figure 1A, of east coast strandings (e.g., six whales at Cruden Bay, Grampian) but generally the pattern is one that reflects the Atlantic margin. On the Irish Coast [Figure 1B] strandings are again pre- dominantly along the western margin, bordering the Atlantic Ocean. Strandings on the coastlines of England and Wales [Figure 1C] are less numerous and mainly occur along the eastern margin (North Sea coast), there have be\en no Sperm Whale strand- ings on the coast of Wales. Strandings on the east Canadian coast are concentrated around the areas of Newfoundland, Nova Scotia and Sable Island. However, it is acknowledged that the low human population densities in the northern part of the conti- nent probably results in substantial under reporting in northern regions of Canada. Data Observations Table 1 summarises annual numbers of animals stranded and the associated number of events during the period 1988 to 1997 for each of the four regions. Figure 2 illustrates these data and several important features can be seen. There have clearly been many more strandings on the Scottish coast than on either the Irish or eastern Canadian coasts, evident by virtue of the fact that all the plots in Figure 2 are on the same vertical scale. There are no visually con- vincing trends, or statistically significant correla- tions, between year and stranding rate (P > 0.2 for each area separately, and for the British Isles as a whole), in the 1988 to 1997 data — i.e., there is nei- ther a clear increase or decrease in strandings with time during this period for any of the regions. Although there are two notable examples of mass stranding in the Scottish data, the trend is generally one of single whale strandings. The English data show single whale strandings in all but one case and the Irish data show only single whale strandings. Multiple strandings are, by contrast, relatively com- mon in eastern Canada (one third of all events). The differences in the proportion of multiple strandings between the three regions (excluding England because of low sample size) are significant (contin- gency-table test, G = 10.76, 2 d.f., P< 0.01). Table 2 summarises the seasonality of strandings during the ten year period 1988 to 1997, and the data are illustrated graphically in Figure 3. The Scottish and eastern Canadian data show peak numbers of THE CANADIAN FIELD-NATURALIST Vol. 116 animals stranding in the autumn and winter, although the trend is less clearly pronounced in the number of events. The Irish data are similar but lack the peak during the October-December quarter. The English data are less numerous but similarly only show events in the Autumn and Winter. Table 3 summarises all documented stranding data for the twentieth century to 1997 as consecutive ten year totals for Scotland, Ireland, England and Nova Scotia (Sable Island). Sable Island is the only Canadian region for which moderately extended records exist, however even for this region data prior to 1987 were collected inconsistently, hence it is not included in further analyses. The data for Scotland, Ireland and England are graphically presented in Figure 4, which shows increases in reported stranding events with time in all regions. The picture is quite striking in the cases of Scotland and Ireland, showing essentially exponential increases in both numbers of whales stranding and the number of stranding events towards the end of the twentieth century. Table 4 summarises the re-worked data as England + Mainland Scotland vs. the more remote Scottish Islands of the Hebrides, Orkney and Shetland. Figure 5 graphically presents the respec- tive trends. Stranding events (and numbers of ani- mals involved) once again show an upward trend for both mainland Britain and the offshore Scottish islands. The Scottish islands account for a greater proportion of stranding events during the most recent decade than during earlier decades, and hence show the greater rate of increase. Numerical Analysis Non-linear regression analysis, using the model of equation (1), yields rates of increase of Sperm Whale stranding events for the whole of the British Isles of approximately 14% per year, beginning at around 1970, which is significantly different from zero (Table 5). There are also substantial, and significant, increasing trends when Scotland, Ireland and England are treated separately. The Scottish rate of increase, 18%/yr, is twice that for Ireland, 9%/yr, and the difference between the two is significant (t- test approximation P<0.05). There are substantially fewer data for England, so estimates are less precise and a significance level cannot be ascribed. There is, however, a significant difference between the strand- ing rates for Mainland Scotland + England vs. the Scottish Islands. The rates of increase stand at 11% /yr from about 1960 for the mainland, and 19%/yr from about 1973 for the Scottish islands. These model results are presented graphically in Figure 6. Stranding events across the British Isles more than doubled during the last two decades of the twentieth century. Discussion There is a general pattern around the British Isles 2002 GOOLD, WHITEHEAD, AND REID: SPERM WHALE STRANDINGS 375 TABLE 1. Strandings of Sperm Whales from 1988-1997 inclusive. Scotland England & Wales Ireland Eastern Canada Year Whales Events Whales Events Whales Events Whales Events 1988 7 i 0 0 4 4 1 1 1989 6 6 0 0 0 0 5 | 1990 4 4 1 1 4 4 6 2 1991 0 0 1 1 0 0 6 3 1992 5 4 0 0 0 0 6 4 1993 3 3 2 2 1 1 0 0 1994 18 8 1 1 1 1 2 2, 1995 4 4 0 0 1 1 1 1 1996 9 4 0 0 1 1 4 4 1997 5 5 3 2 3 3 4 2 TOTAL 61 45 8 7 15 1S SY 21 of Sperm Whale strandings on the western margins bordering the Atlantic Ocean. This might be expect- ed for a species that is adapted to a deep water habi- tat, and which may prefer the submarine slope waters at the edge of the European continental shelf. Data for the most recent ten year period show that there is no Statistically significant trend in the num- Scottish Strandings (1988-1997) 3 E ) z lrish Strandings (1988-1997) 18 j eet a a = Whales CS rr : 2 — B Events 12 ae re a 2 a start a mad Ea) 5 apenas = —- = ss 2 iT | pene . om te S,) 9 A VY S)) s,) s,) a ito ne Ce cal eae s,) Oo) e,) ,) De ee a Year FiGuRE 2. Annual Sperm Whale strandings from 1988-1997. ber of Sperm Whale stranding events on either the British or eastern Canadian coastlines. This observa- tion contrasts strongly with multi-decadal examina- tion of strandings, which reveals a substantial increase in the latter half of the twentieth century across the British Isles as a whole, with Scotland exhibiting the greatest rate of increase. The question English Strandings (1988-1997) 185 16 + @ Whales 12 Number Number 376 THE CANADIAN FIELD-NATURALIST Vol. 116 TABLE 2. Seasonality of Sperm Whale strandings from 1988-1997 inclusive Scotland England & Wales Ireland Eastern Canada Season Whales Events Whales Events Whales Events Whales Events January—March 22 17 1 1 7 v 10 6 April-June 10 10 0 0 3 3 7 6 July—September 9 8 0 0 3 3 6 3 October-December 20 10 7 6 1 1 13 5 TOTAL 61 45 8 7 14 14 36 20 Ireland excludes uncertain 1997 date; Canada excludes 1994 female remains, however, as to why have there been these increases. Some possibilities are discussed here. Increases in Reporting One hypothesis would be that strandings have not increased at all, but that we are seeing an artefact in the data as a result of improved reporting in recent times, perhaps as society becomes more aware of environmental issues and species conservation. If this were indeed the case, one might expect remote outlying regions to have historically been poorly rep- resented in the stranding record, with subsequent increases in reporting as communications and envi- ronmental awareness improve. This hypothesis does seem to be consistent with the fact that the remote Scottish islands exhibit a greater rate of increase in strandings than does the mainland, where communi- cations have perhaps been better during recent histo- ry. However, strandings still have significant expo- nential increases on both the mainland and Irish coasts, suggesting that geographical remoteness from a central recording point is not necessarily a deciding factor. Indeed, one may take the opposing viewpoint and argue that Sperm Whales are large animals and their strandings generate sufficient interest to make reporting via some medium or another highly likely in most instances. If increased reporting effort is sus- pected in recent years, a good control would be to consider the data pre- and post the 1992 onset of the DoE project in Scotland, where an official reporting scheme was put into effect. It has already been demonstrated that no significant trend emerged in the data for the period 1988 to 1997, and inspection of Figure 2 shows no sign of a “surge” in Sperm Whale strandings from 1992. In contrast, strandings of small cetaceans in Scotland show a multi-fold increase at the 1992 project onset. This is just the effect we expect to see with species that are small and less noteworthy of reporting by casual observers. The fact that this does not occur in the Sperm Whale data leads us to suspect that increased reporting is not a major factor in the case of this species. Increases in Population Size Taking the north Atlantic as a whole, and dis- counting effects of differing habitat use, the observed rates of increase in Sperm Whale strand- ings on Irish and Scottish coasts (0.089/yr and 0.181/yr) far outstrip the potential population growth rate according to IWC models of Sperm Whales (0.008/yr) and the possibly more realistic Killer Whale, Orcinus orca, models (0.029/yr) — see Appendix E. Substantial increases in the populations of large males are possible for a few years after sex- biased whaling ends (Figure 7), but they do not appear to last long enough to account for the dramat- ic trends in Scottish and Irish strandings. Furthermore, if the increase in strandings was solely due to an increase in population size, then strandings might be expected to increase at the same rate throughout the population where large males strand. The discrepancy (two fold) between the rates of increase of strandings on Scottish and Irish coasts, both effectively within the same geographic area, suggests that a population increase is unlikely to be responsible for the observed trends. However, differ- ing habitat use, even across a relatively small geo- graphic area, may account for different stranding densities. Defining habitat use over such small scales requires higher resolution data than is currently available, although recent work suggest the Atlantic Frontier adjacent to the Scottish coasts to be an important region for Sperm Whales (Moscrop and Swift 1999). Changes in Distribution and Migration Patterns The temporal changes in British stranding rates could be caused by changes in distribution and migration patterns, but they would have to have been extremely dramatic to fully account for the patterns in the Scottish and Irish strandings. In particular, the waters off Scotland would have had to have changed from an area of very marginal importance before 1975 to a major habitat in the 1990s. Data presented by Evans (1997) are not consistent with this scenario: there were an equal number of sightings of Sperm Whales from the ocean weather ship “Cumulus” 300 km west of Scotland during the peri- ods 1975 to 1984 and 1985 to 1994 (14 each). However, Evans does report an increased preponder- ance of smaller whales stranding since 1975, sug- gesting a shift in the population structure with possi- ble immature animals frequenting higher latitudes. 2002 GOOLD, WHITEHEAD, AND REID: SPERM WHALE STRANDINGS 377 TABLE 3. Sperm Whale strandings within decadal periods from 1908-1997. Scotland England & Wales Ireland Eastern Canada Year Whales Events Whales Events Whales Events Whales Events 1908-1917 ws EPs > 0 > 0 1 1 1918-1927 0 0 0 0 0 0 1928-1937 0 0 ys 2 1 1 1938-1947 3 3 0 0 2 = 1948-1957 1 1 0 0 1 1 1958-1967 1 1 1 1 3 3 1968-1977 4 4 4 4 4 4 >1 =a 1978-1987 9 9 3 3 6 6 ae 3 1988-1997 61 45 8 7 15 15 9 4 TOTAL > 81 > 65 > 18 > iz 33 33 = TS >8 Although there have been many stranded animals < 12 m in length on the British coasts since 1975, there continues to be many strandings of animals > 12 m in length; further there have been incidents. of small whale strandings prior to 1975 — where the data are very much more sparse in any case. Figure 8 illustrates the body length data of Sperm Whales stranded on the Scottish, Irish, English and eastern Scottish Stranding Seasonality @ Whales G Events i : 15 2 i . = 2 10 0 ee bE uo 8 Jan-March April-June July-Sept Oct-Dec Season Irish Stranding Seasonality 25 ] gw Whales 20 @Events | - 15 2 E s 2 10 Jan-March April-June July-Sept Oct-Dec Season Canadian coasts; the wide range of body lengths in recent times (particularly in the Scottish data) is quite evident. Evans (1997) suggests that younger males may have less incentive to periodically return south to the breeding schools than will the large males, and as a consequence may suffer seasonal food shortages, leading to mortality. Post mortem examinations from recent Scottish strandings English Stranding Seasonality 25 4 @ Whales | 20 +— =] G Events i 5 15] fi <2 = = Za 4 Jan-March April-June July-Sept Oct-Dec Season East Canadian Stranding Seasonality 25 | te - | | m™ Whales | | 20 we @ Events i te (0) ame | 2 E ) | Ss / = 10+ —— SE ; : si i. 0 +— + -- +—- +— Jan-March April-June July-Sept Season FIGURE 3. Seasonality in Sperm Whale strandings from 1988-1997. 378 indicate mid-lateral blubber thicknesses of 120 to 140 mm, suggesting that these animals are in a rea- sonably good nutritive state. Given the continued Decadal Scottish Strandings 7 eerste 25: 0 @ Whales Sarid Bid G Events 50 +-— - ——— - ~ 5 AG — : 2 = 2 30;— sos a 20 + —— 10 i col ah molt FL age IE AN AO gh Py RO ce bE he Behe | Pie? ©) Re) Re) Re) Re Re Re) Re) Re Decade Decadal English Strandings 8 - : @ Whales G Events o 2 £ =) z ay % rn vod ov Decade Decadal Irish Strandings i SE pe | @ Whales | 14 L-—. 12 ce 3 E 8 s 2 6. 4 2 eee: a ae wi wv we a @ ow eo oO A idee ca ae eile ea Decade FIGURE 4. Decadal trends in Sperm Whale strandings from 1908-1997. THE CANADIAN FIELD-NATURALIST Vol. 116 stranding of males > 12 m in length, and the reason- able body condition of many stranded animals, Evans’ argument is not entirely convincing. Increased Mortality Fisheries By-Catch Fisheries by-catch is a serious problem around the British coasts for a number of small cetacean species (Kuiken et al. 1994; Kirkwood et al. 1997; Berrow and Rogan 1998). Harbour porpoise are thought to be suffering a population mortality rate of some 6% per year due to bycatch, and many beached carcasses show signs of entanglement in fishing gear. Common dolphin have suffered similar instances of entanglement, with notable mass stranding of car- casses where fishing injury has been involved (Kuiken et al. 1994). Although fisheries and driftnet- ting has been a serious problem for Sperm Whales in some regions of the world [e.g., the Mediterranean (Di Natale 1990)] there is no gross external evidence from recent post-mortem examinations of British and eastern Canadian stranded Sperm Whales having suffered entanglement in fishing gear. Historical records similarly do not suggest fisheries interaction to be a factor in strandings. We therefore consider the increasing rate of Sperm Whale strandings unre- lated to direct fishing injury. Further, the prey species of Sperm Whales are typically dissimilar to those harvested commercially for human consump- tion (Clarke 1996), so direct conflict with fishery resources also seems unlikely. Nutrition & Toxic contamination It is difficult to perform even a basic autopsy in the remote locations where Sperm Whales often strand. Conversely, public health concerns often restrict beach-head autopsies in more populated regions. Information on the health status of stranded Sperm Whales is therefore rather limited. However, stomachs from whales which have been examined, particularly from two mass strandings in Scotland (Santos 1998) generally show evidence of recent feeding, i.e. squid beaks are present — mainly Gonatus fabricii. In addition, mid-lateral blubber thicknesses (of some 120 to 140 mm) suggest whales that strand on the Scottish coast to be in a reasonable nutritive state prior to death, tending to weigh against chronic disease as a cause of stranding. These obser- vations contrast with those of recent Sperm Whale strandings on the Belgian and Dutch coasts, where whales had reduced blubber thickness and some had severe weight deficiency (Jauniaux et al. 1997). This may be consistent with the notion of the North Sea (to the east of mainland Britian) acting as a Sperm Whale trap (Smeenk 1997), resulting in starvation as this shallow shelf sea is largely devoid of typical Sperm Whale prey items. However, our data mainly considers strandings on the western coasts of the British Isles, and starvation is not evident. 2002 GOOLD, WHITEHEAD, AND REID: SPERM WHALE STRANDINGS 379 TABLE 4. British strandings data tabulated as mainland (excluding Ireland) vs. remote islands. England + Mainland Scotland Year Whales 1913-1917 1918-1927 1928-1937 1938-1947 1948-1957 1958-1967 1968-1977 1978-1987 1988-1997 Total oe ey) OS ch eeaeeeoeeeata Considering the North Sea again, Mercury and Cadmium were also found at high levels in Sperm Whales stranded on the Belgian coast (Bouquegneau et al. 1997). Cadmium, although expected due to a cephalopod rich diet, was found at concentrations twice that previously recorded in the literature, and was not bound to metallothioneins (a protein known for its protective effect against heavy metals). Measured chlorobiphenol levels in British stranded Sperm Whales have been shown to be low in com- parison with other species of cetacean (Wells et al. 1997). Chemical pollution is therefore a factor for consideration in Sperm Whale strandings generally, but given the relatively good condition of recent whales stranded on the Scottish coast, it is unlikely that toxic contamination is a direct cause of such dramatic increases in strandings as are observed in the data. Industrial Activites Other than fishing, which has been considered sep- arately, the anthropogenic activities most likely to impact the deep water environment of the Sperm Whale are shipping traffic, oil and gas exploration, England & Mainland Scotland Number ar oe wr er ae SL a Le fate nner Meee lke” a IR oll aR MN a Se Decade Events Nm] Sco aes eae Scottish Islands Whales Events Q |S BlonNnpRrReR WOO SC re [ WOIlrNInNAN BRR WOC? and military activities. Sperm Whales are deep water pelagic predators, feeding primarily on water column species and also some benthic species. Therefore, they are likely to be directly affected by activities that impact on the entire water column in which they live. All three of the above mentioned activities entail varying degrees of physical and acoustic intrusion into the water column. Considering the case of shipping, the passage of a vessel through the water presents a possible collision hazard to a whale at the surface (Terhune and Verboom 1999). Further, the noise emitted from engines and associated machin- ery serves to raise the levels of background noise, to which Sperm Whales may or may not be sensitive. Further, raised levels of background noise may pro- gressively mask sounds of importance to sperm whales — e.g., intra-specific communications; biosonar for navigation and food finding; sounds created by other marine organisms. Large powered vessels have been traversing the Atlantic since the beginning of the twentieth century, but the rapid increase in strandings is a relatively recent phe- nomenon. Although fast vessels present a serious Scottish Islands 50 - oe a ae 40 +— GSeries2 a Re eee eee Pe eee ae ee 30 + Number N oi FIGURE 5. Decadal Sperm Whale stranding trends separated into mainland Scotland + England vs. more remote Scottish Islands. 380 THE CANADIAN FIELD-NATURALIST Vol. 116 - TABLE 5. Parameter estimates for the model of increase in Sperm Whale stranding events by year. Standard errors are indi- cated in parentheses. Geographic Area Basal level [k] /yr British Isles Sok (3) Scotland 1.1 (0.6) Ireland 1.0 (0.5) England 0.2 (0.8) England +Mainland Scotland 0.5 (0.6) Scottish Islands 1.2 (0.6) hazard to Sperm Whales at the surface, and multiple mortalities have resulted from fast ferry strikes in other regions of the world (Aguilar et al. 2000), such vessels do not operate intensively in the regions under consideration here, and there is no evidence of vessel collision from the strandings data. S Stranding Events 8 S) 10F- 1920 1930 1940 1950 1960 1970 1980 1990 2000 Year 8 Ai Stranding Events & 8 1950 1960 1970 1980 1990 2000 Year {e200 1930-=—«1940 FIGURE 6. Fitted curves of exponential growth showing (a), Sperm Whale stranding trends for Scotland, Ireland, England and the British Isles as a whole; and (b) mainland Scotland+England vs. remote Scottish Islands. Increase [b] /yr Start of increase [c] L-139"(Q.0T9) 1970 (6) 1.180 (0.019) 1971 aates 1.089 (0.016) 1962 (10) 1.058 (0.027) 1928 (102) 1.105 (0.024) 1960 (17) 1.188 (0.028) 1973 ia) Oil and gas exploration has been demonstrated in some instances to cause behavioural reactions in large whale species (Richardson et al. 1995), and avoidance of high powered sound sources by Sperm Whales has been documented (Bowles et al. 1994). The type of activity of primary concern is seismic surveying, where loud, concussive type sound pulses are produced by high pressure air-guns beneath the water surface. Seabed sub-bottom profiles are mapped through the process of penetration and reflection of these sound pulses, which are typically received by long trailing hydrophone streamers deployed from seismic vessels. Seismic pulse energy also radiates outwards into the wider water column, essentially ensonifying a large swath of ocean. Behavioural avoidance of these activities has been documented in Bowhead, Gray and Humpback whales (Richardson et al. 1995; Richardson 1998; McCauley 1998, 2000) and there is some evidence that sperm whales may be displaced by such activi- ties (Mate et al. 1994). In addition, there is evidence that seismic pulses also have the potential to cause disturbance to small odontocetes (Goold and Fish 1998), which should be the least vulnerable of all cetaceans to these noises. Seismic surveying has Proportional increase in males per year 0 5 10 15 20 Year FIGURE 7. Rates of increase of older (> 20yr old) males, following the end of exploitation which resulted in the proportion of mature males in the population being artificially reduced by 80%, 60%, 40%, 20% or 0%. 2002 Scotland — Body Lengths et ' ' ' ‘ ' ‘ ‘ t 4 ' ' ' ' ' ) ‘ r ' ' ' ' ‘ ‘ ' . ' ' ' ' ' ' ' ‘ cr ‘ ‘ ' ‘ ‘ ' ' 25 i 1 ' ' ‘ ' ' c ' ' ‘ ‘ ' ' ' ' cr ‘ ' '‘ ' 1! ' ' _ Body Length (m) asin a om Sitti: teliteielitel itt. tt lteter rs ltt tt ns lteitietets teliietet f i ts meee eens 0 1900 1920 1940 1960 Year 1980 2000 2020 GOOLD, WHITEHEAD, AND REID: SPERM WHALE STRANDINGS 381 lreland — Body Lengths ' c ' ‘ 1‘ ' ' ‘ ‘ ' r i ' '‘ ' ‘ ' ce) Body Length (m) i ee 2020 25 20 Body Length (m) 0 1900 1920 1940 1960 1980 Year FIGURE 8. Body lengths of Sperm Whales stranded on the British and eastern Canadian coastlines. been conducted for decades and there are conflicting reports as to its effects on marine wildlife. However, it is notable that extensive deep water seismic sur- veying took place during the 1990s in the Atlantic Frontier region, to the west of Scotland, approxi- mately adjacent to the remote Scottish Islands where the highest strandings rate has occurred. However, the Scottish Sperm Whale stranding data show peaks during the autumn and winter, whereas seismic oper- ations tend to concentrate during the summer months in the Atlantic Frontier. The same scenario of sea- sonal mismatching is true for Nova Scotia too. These data, therefore, do not support a hypothesis of cause and effect between strandings and seismic explo- ration, unless there is some kind of progressive audi- tory fatigue/damage, resulting in disorientation and a lead time to stranding events. Data on both Sperm Whale abundance and line miles of seismic shooting in either region are too sparse to draw any correla- tions. Physiological evidence of acoustic trauma in carcasses is also very cryptic, requiring detailed examination of ears and other tissue structures, which is currently beyond the scope of the stranding schemes. Military Activities Military activities are by their nature classified, and hence data are very sparse. Potential impacts again include vessel strikes and/or acoustic distur- bance by high powered sonar systems and ordnance detonation. However, as with seismic surveying, there is as yet no evidence to suggest that military activities have played a direct role in Sperm Whale 382 strandings, although there is evidence from other regions of the world that mid-frequency military sonars can cause mass strandings of deep diving whales (Frantzis 1998). The best documented inci- dent was the multiple species strandings in the Bahamas during March 2000. A number of deep diving beaked whales, along with two Minke Whales and a dolphin, stranded alive in a highly atypical manner, but with timing and position highly correlat- ed with naval activity and the deployment of high powered, tactical mid range sonar systems. Post mortem analysis found haemorrhaging in the brain and ear tissue, consistent with acoustically induced trauma (Balcomb and Claridge 2000; NMFS 2001). Conclusion There has been a large increase in reported Sperm Whale stranding events on the British coast during the latter half of the twentieth century. Available data suggest that this phenomenon is not mirrored on east Canadian coasts, although the available data are too sparse to lend weight to this suggestion. British strandings have increased most rapidly off the Scottish coast, with most of the increase due to strandings in the outer islands of Hebrides, Orkney and Shetland since about 1973. There is no single satisfactory explanation, but the data do not support the increase in strandings as being a simple result of an increase in population size. Instead, it seems like- ly that the trend has two or more causes which may be acting synergistically. Valid hypotheses include increased reporting and anthropogenic effects. Cause(s) and effect(s) appear to be manifested in greatest concentration near the Scottish Islands. Acknowledgments Thanks to Zoe Lucas and Sascha Hooker for pro- viding Sable Island data, Emer Rogan and Mick Mackey for providing the Irish data, Alex Muir for providing the English data and Rod Penrose for con- firming zero strandings in Wales. We acknowledge the DETR (formerly DoE) for financial support of the strandings projects in the UK. Thanks to HM Coastguard and the Receiver of Wrecks for allowing access to carcasses for necropsy. 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Life his- tory and population dynamics of resident killer whales (Orcinus orca) in the coastal waters of British Columbia and Washington State. Reports of the International Whaling Commission (special issue), 12: 209-243. Reeves, R. R., and H. Whitehead. 1997. Status of the sperm whale, Physeter macrocephalus, in Canada. Canadian Field Naturalist 111: 293-307. Richardson, W. J., C. R. Greene, C. I. Malme, and D. H. Thompson. 1995. Marine mammals and noise. Academic Press. Richardson, W. J. 1998. Reactions of Bowhead whales and Ringed seals to an open water seismic program in the Alaskan Beaufort Sea. In Workshop Documentation, Seismic & Marine Mammals Workshop, 23-25 June, London. Santos, M. B. 1998. Feeding ecology of harbour porpois- es, common and bottlenose dolphins and sperm whales in the northeast Atlantic. PhD thesis, University of Aberdeen. Sheldrick, M. C. 1989. Stranded whale records for the entire British Coastline, 1967-1986. Pages 298-329 in Investigations on Cetacea, Volume XXII, Edited by G. Pilleri. Sheldrick, M. C., P. J. Chimonides, A. I. Muir, J. D. George, R. J. Reid, T. Kuiken, C. Iskjaer-Ackley, and A. Kitchner. 1994. Stranded cetacean records for England, Scotland and Wales, 1987-1992. Pages 259-283 in Investigations on Cetacean, Volume XXV. Edited by G. Pilleri. Smeenk, C. 1997. Strandings of sperm whales, Physeter macrocephalus, in the North Sea: history and patterns. Bulletin de L’ Institut Royal des Sciences Naturelles de Belgique, Biologie, Supplement, 67: 15-28. Strong, D. in press. Sperm whale (Physeter macrocephalus L.). Irish Naturalist’s Journal. Terhune, J. M., and W. C. Verboom. 1999. Right whales and ship noises. Marine Mammal Science 15(1): 256-258. Wells, D. E., C. McKenzie, and H. M. Ross. 1997. Patterns of organic contaminants in marine mammals with reference to sperm whale strandings. Bulletin de L’Institut Royal des Sciences Naturelles de Belgique, Biologie, Supplement, 67: 91-103. Whitehead, H. 1995. Status of Pacific sperm whale stocks before modern whaling. Reports of the International Whaling Commission 45: 407-412. Received 10 August 2000 Accepted 28 June 2002 384 THE CANADIAN FIELD-NATURALIST Vol. 116 Appendix A: Scottish Strandings Raw Data, 1913-1997 Reference Date Sex Length (m) Location of Stranding Harmer (1927) 18 Dec 1913 14.63 Fort George, Inverness ‘4 23 May 1917 M 18.08 Latheron, Caithness Fraser (1953) 07 Dec 1938 Hebrides : Sep? 1946 M c60ft? Shetland : Oct? 1946 M 14.94 Shetland Fraser (1974) 30 Jun 1955 8.23 Bay Head, North Uist, Hebrides 4 ?? Jun 1958 c18.29 North Roe, Shetland Scheldrick (1989) ?? Jan 1975 M 10.06 Holm of Boray, Isle of Gairsay, Orkney Islands : 12 Sep 1975 M 13.41 North Boisdale, South Uist, Outer Hebrides a 01 Jul 1977 M eee North Boisdale, South Uist, Outer Hebrides a 20 Nov 1977 M 18.29 Ham Voe, Foula, Shetland Islands “ 28 Feb 1979 M c18.29 Tinna’s Voe, Skeld, Shetland Islands . 05 Mar 1979 M 16.76 West Gernish, South Uist, Western Isles s 22 Aug 1979 M 14.12 Cullen Bay, Grampian af ?? Jan 1983 M c11.58 Flotta Island (S.E.), Weisdale Voe, Shetland - ?? Jun 1983 M 7.62 Isle of Tiree, Strathclyde ‘ ?? Jun 1984 M c9.14 Gortanaoid Point, Islay, Strathclyde ‘ 13 Dec 1984 M POT Bea Ness, Sanday, Orkney Islands i 23 Jan 1985 M Crouie, Gardenstown, Grampian : e 04 Feb 1986 10.67—12.19 Drid Geo (S.Side), Eshaness, Mainland, Shetland Scheldrick et al. (1994) v 27 Feb 1988 M 115 Galoson, west coast Isle of Lewis, Western Isles é 26 Feb 1988 M ey 2 Sanday, Unst, Orkney Islands 2 19 Mar 1988 M 14 Bay of Newark, Orkney Islands 24 Mar 1988 M 18.29 Lendalfoot, (nr Girvan) Strathclyde A 25 Mar 1988 12.19 Garrynamonie, South Uist, Western Isles 4 07 Jul 1988 M Bressay, Shetland Isles 4 29 Nov 1988 M 11 Bow Head, Westray, Orkney Islands f 22 Feb 1989 18.3-21.3 Culswick West, Shetland Isles 16 May 1989 9.14 Isle of Eigg, Highland % 15 Jun 1989 M 12.19 The Ness, North Mainland, Shetland Isles | “ 27 Jul 1989 M 14.33 North Beach, Haroldswick, Unst, Shetland Isles | ‘ 01 Aug 1989 M 10.67 Heagan Beach, West Burrafirth, Shetland Isles ‘ 02 Aug 1989 M 18.29 Stoney Bridge, South Uist, Western Isles | ‘ O01 Feb 1990 M 14.94 The Bar, Culbin Forest, Findhorn, Grampian . ‘! 23 Feb 1990 M 14.02 Daliburgh, South Uist, Western Isles , 04 Apr 1990 10.67 Cragaig, Isle of Ulva, Strathclyde : ‘: 04 Apr 1990 M 11.8 Rubh An T Subhein, (nr Rendle) Isle of Mull # 29 May 1992 M c 16 Fuday, Barra, Western Isles is 20 Jul 1992 M Monach Isles, Western Isles H ‘ 20 Jul 1992 Monach Isles, Western Isles ¢ 04 Dec 1992 M c 10 Sanday, Orkney Islands | * 22 Dec 1992 Northton, Harris, Western Isles | Reid 21 Jul 1993 The Burr, Eshaness M 21 Nov 1993 M hey Lochailort, Highland* | ‘ 24 Nov 1993 M 15.65 Loch Ainort, Skye* ! ” 13 Mar 1994 M 13 Hougharry, North Uist, Western Isles " 13 Mar 1994 M LOT Ayre of Birrier, Yell, Shetland Isles < 27 Mar 1994 M 16 Campa Islay, Argyll 5 09 Apr 1994 cl6 Sound of Berneray, Western Isles i ‘ 24 Apr 1994 M cl0 St. Ninians Isle, Shetland | ‘ 30 May 1994 cl2 Floating off Balta, Shetland N 18 Aug 1994 Swordle, Ardnamurchan, Highland - 07 Dec 1994 M 12.4 Backaskail Bay, Sanday, Orkney Islands* | ¢ 07 Dec 1994 M 12.3 Backaskail Bay, Sanday, Orkney Islands* | 07 Dec 1994 M 12.8 Backaskail Bay, Sanday, Orkney Islands* | F 07 Dec 1994 M 12.8 Backaskail Bay, Sanday, Orkney Islands* ‘ 07 Dec 1994 M ey Backaskail Bay, Sanday, Orkney Islands* } ‘ 07 Dec 1994 M 12.6 Backaskail Bay, Sanday, Orkney Islands* | i 07 Dec 1994 M 12 Backaskail Bay, Sanday, Orkney Islands* continued 2002 GOOLD, WHITEHEAD, AND REID: SPERM WHALE STRANDINGS 385 Appendix A: continued Reference Date 07 Dec 1994 07 Dec 1994 07 Dec 1994 07 Dec 1994 19 Jan 1995 27 Jan 1995 23 Mar 1995 09 Apr 1995 28 Jan 1996 28 Jan 1996 28 Jan 1996 28 Jan 1996 28 Jan 1996 28 Jan 1996 08 Sep 1996 27 Dec 1996 31 Dec 1996 28 Mar 1997 31 Mar 1997 07 Apr 1997 20 Nov 1997 27 Nov 1997 Italics = mainland strandings * = strandings in which animals are known to have come ashore alive N x ss ex [ssh ccs SSoeee e Length (m) 13.3 12.4 125 13.4 cl6 cl4 13.7 cll 13.65 TRGS 12.6 W375 i271 12.85 14.4 c13.9 el3 75.2 c13 14.17 15.54 Location of Stranding Backaskail Bay, Sanday, Orkney Islands* Backaskail Bay, Sanday, Orkney Islands* Backaskail Bay, Sanday, Orkney Islands* Backaskail Bay, Sanday, Orkney Islands* Brenish Beach, Lewis, Western Isles Culla Beach, Benbecula, Western Isles Nr Carse of Ardersier, Highland* Creag Loisgte, Islay, Strathclyde Cruden Bay, Grampian* Cruden Bay, Grampian* Cruden Bay, Grampian* Cruden Bay, Grampian* Cruden Bay, Grampian* Cruden Bay, Grampian* Largelsland of Mousa, Shetland Traigh Angus, Islay, Strathclyde Sand, West Mainland, Shetland Goile Chroic, Lewis, Western Isles Airth, Central* Papa Stronsay, Orkney Islands Stornoway Airport, Western Isles* Melbost, Western Isles* Appendix B: England and Wales Raw Data, 1913-1997 Reference Fraser (1946) Sheldrick (1989) Sheldrick et al. (1994) Muir Date 18 Jan 1934 25 Jan 1937 2? Nov 1967 07 Feb 1973 06 Oct 1973 19 Jan 1974 18 Nov 1977 17 Sep 1980 01 Mar 1985 30 Nov 1986 09 Feb 1990 12 Nov 1991 02 Dec 1993 15 Dec 1993 10 Nov 1994 03 Dec 1997 03 Dec 1997 03 Dec 1997 Sex M M xs Sano oe oS. Se Length (m) 10.29 18 15.54 c41ft 14.94 13.41 5.94 40—5Oft 15.32 Wa 72 ES 12.5 15.6 15:85 14.4 c12.19 c12.19 Location of Stranding Kimmeridge, Dorset Bridlington, Yorkshire _ Samson, Isles of Scilly, Cornwall Portreach, Cornwall Seaton Point, Boulmer, Northumberland Skidbrooke, Lincolnshire Boule Bay, N.coast Jersey, Channel Isles Marazion, Cornwall Skegness, south of Lincolnshire Holcombe Beach, Wells, Norfolk Long Rock Beach, Marazion, Cornwall Brancaster, Norfolk Hunstanton, Norfolk Atwick, Humberside Hawkster Bottoms, nr Whitby, North Yorkshire Skegness, Lincolnshire Trinity Sand, Humber Estuary Trinity Sand, Humber Estuary 386 THE CANADIAN FIELD-NATURALIST Appendix C: Irish Strandings Raw Data, 1901-1997 Vol. 116 Reference Date Sex Length (m) Location of Stranding Berrow and Rogan (1997) 04 Sep 1916 5.49 Roundstone, Galway 4 15 Jul 1936 M Lislary, Sligo @ 18 Nov 1939 Kilkee, Clare f 22: 277 1943 Louisberg, Mayo (L7477) ‘ 02 Jan 1952 M pp Derryloughan, Galway if 25 Jul 1963 M 20 Rosbeg, Donegal i 27.277 1967 Bertraghboy Bay, Galway (L7437) ( 21 Mar 1967 Achill Isle, Mayo i 2? Dec 1968 c9-12 Crookhaven, Cork y 20 May 1972 c12-15 Buncrana, Donegal i 30 Nov 1974 M 13 Portrush, Antrim (C8541) 09 Jul 1977 M 14.2 Sligo (G6139) J 13 Oct 1978 Kerry (Q7424) i ?? Jan 1980 Rathlin Isl. Antrim (D1050) ¢ 2? Sep 1980 Bantry Bay, Cork (V8241) . 2? Jul 1982 10.7-12.2 Kerry (V3298) j 08 May 1987 Maglilligan St., Co. Londonderry (C6738) f 01 Jan 1987 M 20 Kerry (V4898) +f 19 Mar 1988 M 12.65 Clare (R0381) ' 13 Apr 1988 M Mayo i 15 Jun 1988 M 9 Mayo (G1141) 09 Aug 1988 M fife 38) Sligo (G5941) i 07 Feb 1990 M 13.7 Mulranny, Mayo (L8296) 4 16 Mar 1990 M Carrickfin, Donegal (B7923) : 30 Mar 1990 M Portscaman, Donegal (C4156) J 10 Oct 1990 Minard, Kerry (Q5298) Mallarney and 26 Sep 1993 F 11.07 Kilmore Quay, Co. Wexford (S9603) Smiddy (unpublished) Berrow and 16 Feb 1994 M 10.4 Inverin, Co. Galway (M0521) Rogan (1997) Berrow and 15 Jun 1995 F 122 Youghal, Co. Cork (C0874) Rogan (1997) Gassner and 29 Mar 1996 M 14.8 Tory Island, Co. Donegal (B8546) Gatins (unpublished) Kennedy (in press) ?? Mar 1997 M c40ft Maghery, Co. Donegal (B7110) IWDG 20 Sep 1997 c40ft Oranmore, Co. Galway (M3824) Strong Sept/Oct 1997 6~7 Blacksod, Co. Mayo (F6418) Appendix D: Eastern Canadian Strandings Raw Data: ~1987-1997 (with Sable Island from 1970-1997) Reference Date Sex Length (m) Location of Stranding Lucas and Hooker (2000) 2? Apr 1977 Sable Island, Nova Scotia Lien (personal communication) 8 Jun 1979 F 13.4 Burnside, Bonavista Bay, Newfoundland/Labrador : 31 Aug 1979 M 6.9 Portugal Cove, Conception Bay, Newfoundland/Labrador ; 14 Aug 1981 M 16.1 Reefs Harbour, Newfoundland/Labrador ‘ 5 Oct 1981 M Fe Nain, Newfoundland/Labrador Z 10 Apr 1982 M 10.0 Lords Cove, Newfoundland/Labrador i 28 Apr 1982 M 14.4» Point May, Newfoundland/Labrador . 29 Jun 1982 M 13.2 Cartwright, Newfoundland/Labrador i. 25 Jul 1983 M 15.0 Hare Harbour, Newfoundland/Labrador continued 2002 GOOLD, WHITEHEAD, AND REID: SPERM WHALE STRANDINGS 387 Appendix D: continued Reference Lucas and Hooker (2000) ” Reeves and Whitehead (1997) ” Lien (personal communication) Reeves and Whitehead (1997) Lucas and Hooker (2000) Reeves and Whitehead (1997) Lien (personal communication) Whitehead (unpublished) Lien (personal communication) Lucas and Hooker (2000) Lien (personal communication) Date 2? Jul 1983 ?? Feb 1984 ?? Mar 1987 11 Jun 1987 08 Jul 1987 13 Dec 1988 01 Mar 1989 01 Oct 1989 01 Oct 1989 01 Oct 1989 01 Oct 1989 01 Oct 1989 01 Oct 1989 30 Sep 1990 30 Sep 1990 30 Sep 1990 30 Sep 1990 06 Oct 1990 06 Oct 1990 15 Jan 1991 25 Apr 1991 25 Apr 1991 19 Dec 1991 20 Dec 1991 23 Dec 1991 17 Jan 1992 17 Jan 1992 17 Jan 1992 ?? Mar 1992 13 Jun 1992 24 Jun 1992 ?? Jul 1994 2? Dec 1994 06 Mar 1995 10 May 1996 ?? Jun 1996 3 Jul 1996 20 Jul 1996 13 Jan 1997 13 Jan 1997 13 Jan 1997 17 July 1997 Sex Sed a MES £ SESSSSSSSSE SSS E5 as & SSS SS M Length (m) >6.0 16.5 13.6 14.0 14.5 14.6 4 13.1 cl5 12.0 2 12.8 31 11.55 12.0 Location of Stranding Sable Island, Nova Scotia Sable Island, Nova Scotia Sable Island, Nova Scotia Red Harbour, Placentia Bay, Newfoundland/Labrador Penguin Island West, South Coast, Newfoundland/Labrador Near Nail Pond, N.W., Prince Edward Island Conception Harbour, Conception Bay, Newfoundland/Labrador Covehead Harbour, N., Prince Edward Island* Covehead Harbour, N., Prince Edward Island* Covehead Harbour, N., Prince Edward Island* Covehead Harbour, N., Prince Edward Island* Covehead Harbour, N., Prince Edward Island* Covehead Harbour, N., Prince Edward Island* Black Tickle, Labrador* Black Tickle, Labrador* Black Tickle, Labrador* Black Tickle, Labrador* Sable Island, Nova Scotia Sable Island, Nova Scotia Port aux Choix, West Coast, Newfoundland/Labrador Isle Petitgras, Cape Breton Island, Nova Scotia Isle Petitgras, Cape Breton Island, Nova Scotia North Rustico, N., Prince Edward Island Brackley Beach, N., Prince Edward Island Brackley Beach, Prince Edward Island Sable Island, Nova Scotia Sable Island, Nova Scotia Sable Island, Nova Scotia Sable Island, Nova Scotia Port Meunier, Anticosti Island, Quebec Sheldrake 50°15'N, 64954’W, Quebec Cherry Hill, S. Shore, Nova Scotia River John, N. Coast, Nova Scotia Lord’s Cove, Burin Peninsula, Newfoundland/Labrador Burgeo, Newfoundland/Labrador N.W. Miquelon, St. Pierre et Miquelon Grand Bank, Newfoundland/Labrador Burgeo, Newfoundland/Labrador Sable Island, Nova Scotia* Sable Island, Nova Scotia* Sable Island, Nova Scotia* Grand Beach, Fortune Bay, Newfoundland/Labrador *strandings in which animals are known to have come ashore alive Excludes 5 and 6 October 1989 Prince Edward Island Strandings, as probably included in | October total 388 THE CANADIAN FIELD-NATURALIST Vol. 116 Appendix E. Potential Rates of Sperm Whale Population Increase Using the most recent set of Sperm Whale popula- tion parameters agreed to by the Scientific Committee of the International Whaling Commission (age of first birth of female = 11 yr; maximum birth rate of mature females = 0.25 /yr; mortality of females = 0.055 /yr; mortality of males = 0.066 /yr; mortality of juveniles = 0.0926 /yr; equal sex ratio at birth; International Whaling Commission, 1982) Sperm Whale populations with a stable age distribu- tion can increase at a maximal rate of 0.0082 /yr (Whitehead 1995). The IWC Sperm Whale model parameters are not especially reliable (Christal 1998). For instance, fecundity rates of female Sperm Whales seem to be age-specific (Best et al., 1984), and assumed mortality rates seem high for adults, and low for juveniles, when compared with those estimated more recently for other odontocete populations (e.g. Olesiuk et al. 1990). The most reliable life history parameters available for any large odontocete are those for the Killer Whales off Vancouver Island (Olesiuk et al. 1990). These animals have a maxi- mum potential rate of increase of 0.029 /yr with a sta- ble age distribution, and this may be a better estimate for Sperm Whales than the 0.0082 /yr calculated from the assumed Sperm Whale population parameters (Christal 1998), although as Killer Whales prey on Sperm Whales, but Killer Whales themselves have few if any natural predators, the sperm whale rate may be somewhat lower. Recent sperm whaling has concentrated on large males, altering the sex ratio. Thus it is likely that proportions of large males were artificially low when whaling ceased. This means that, following the end of whaling, the population of large males can increase more quickly than that of the females for some years while the sex ratio is imbalanced. This is shown in Figure 8 for rates of increase of popula- tions of large males (> 20 yrs old), whose represen- tation in the population had been artificially decreased, using the population parameters of the IWC (as given above) and assuming stable age dis- tributions of females and young males. For 5—10 years following artificial reduction of the sex ratio, the population of mature males can increase at rates above 0.05 /yr, but over longer periods the effect is small (Figure 7). Bat Night Roost at an Abandoned Mine in Western Maryland SALVATORE J. AGOSTA!, KELLIE M. KUHN?, and DAvID MORTON Department of Biology, Frostburg State University, Frostburg, Maryland 21532-1099 USA Present address: Department of Biology, Leidy Laboratories, University of Pennsylvania, Philadelphia, Pennsylvania 19104 USA.; Corresponding author: agosta@sas.upenn.edu 2Present address: Department of Biology, University of Nevada, Reno, Nevada 89557 USA Agosta, Salvatore J., Kellie M. Kuhn, and David Morton. 2002. Bat night roost at an abandoned mine in western Maryland. Canadian Field-Naturalist 116(3): 389-392. Few published records of night roosts used by bats in eastern North America exist. We describe seasonal use and species composition at an abandoned mine used as a night roost by bats in western Maryland. A total of 81 bats from five species were captured from July to September 1999 and March to September 2000. Big Brown Bats (Eptesicus fuscus) used the mine regularly and were the most abundant species captured. Capture success (number of bats / trap hour) was positively correlated with nightly ambient temperature, but not correlated with relative humidity. Key Words: bats, night roost, mine, Eptesicus fuscus, Pipistrellus subflavus, Myotis lucifugus, Myotis leibii, Lasiurus bore- alis, Maryland. Temperate-zone bats roost in a variety of struc- tures, each important for survival and reproductive success (Kunz 1982). In addition to maternity roosts, summer day roosts, and winter hibernacula, many bats use night roosts in close proximity to foraging areas (Hirshfeld et al. 1977; O’Shea and Vaughan 1977; Kunz 1982; Lewis 1994; Perlmeter 1996; Pierson et al. 1996; Adam and Hayes 2000). For temperate-zone bats, night roosts probably function primarily as resting places that facilitate digestion between nightly foraging bouts (Hirshfeld et al. 1977; Pierson et al. 1996). Although temperate bats spend a high proportion of time in night roosts (e.g., Anthony et al. 1981; Barclay 1982), information on night roost location, activity patterns and species composition is sparse (Adam and Hayes 2000). Recent studies in Oregon (Perlmeter 1996; Adam and Hayes 2000) and California (Pierson et al. 1996) have drawn attention to the importance of man-made structures as night roosts for a variety of species of bats. These studies have emphasized the need for further investigation into night-roosting behavior and night roost selection. Other than a study on the night- roosting behavior of Little Brown Bats (Myotis lucifugus) in an abandoned New Hampshire barn (Anthony et al. 1981), few definitive records of bat night roosts in eastern North America exist. In this paper we report data on the species composition and seasonal use of an abandoned mine used as a night roost by bats in western Maryland. To our knowl- edge, this is the first record of a site used specifically as a night roost by bats in Maryland. Methods We studied night roost use by bats at Round Top Mountain (RTM) located along the Potomac River, 5.4 km southwest of Hancock, Washington County, Maryland (39°N, 78°W). Previous work at a series of abandoned mine tunnels at RTM (Gates et al. 1984; Marsh 1998; D. Feller, personal communication) indicated that these mines serve as important winter and summer habitat for a number of species of bats. Consequently, access to the mines was restricted with gates in 1998 (D. Keech, personal communication). As part of a larger study (Agosta 2001) on the feeding ecology of the Big Brown Bat (Eptesicus fuscus), we mist-netted the entrance of RTM mine Number 3 once every 7-10 days from July to September 1999, and from March to September 2000. Mist nets were always opened about 30 min after sunset. Nets were left open for an average of 3.7 + 0.9h (n= 28 nights of trapping), excluding one night when nets were closed after 45 min because of rain. Captured bats were held in styrofoam cups with perforated lids until the end of each trapping period to eliminate the possi- bility of recapture. To examine possible environmental factors that could influence bat use of the night roost we mea- sured nightly ambient temperature and relative humidity throughout the 2000 study period. Ambient temperature was measured along the forest access road adjacent to the mine three times each night (on the hour, beginning approximately at sunset) with a pocket thermometer. Relative humidity was mea- sured simultaneously with a wet bulb-dry bulb sling psychrometer. Results and Discussion To determine if bats roosted in the mine during daytime, we always watched the mine entrance for activity beginning | h before sunset until it was too dark to observe. Although bats were regularly caught 389 390 THE CANADIAN FIELD-NATURALIST Vol. 116 TABLE 1. Sex composition, age structure and capture times for bats caught at the abandoned mine in 1999 and 2000. Species Male Female Eptesicus fuscus 33 6 Pipistrellus subflavus 13 3 Myotis lucifugus 3 1 Myotis leibii ] Lasiurus borealis 1 aall juveniles were males caught in August 2000 at the mine entrance, we never observed bats exiting the roost for their first foraging bout on any nights in 1999 (n= 6) or 2000 (n = 22). In addition, no bats were seen roosting on two daytime trips into the rel- atively shallow mine (<50 m in length). For each species, capture times were consistent with arrival at the mine following the first foraging bout (Table 1). Finally, we observed numerous bats avoiding our net and entering the mine during periods when other bats were captured. Taken together, these observations indicated that mine Number 3 was used exclusively as a night roost. A total of 81 bats from five species were captured at the mine entrance (1999, n = 17; 2000, n = 64; Table 1). These included the Big Brown Bat (1999, n= 11; 2000, n= 36), Eastern Pipistrelle (Pipistrel- lus subflavus: 1999, n = 2; 2000, n = 21), Little Brown Bat (1999, n = 4; 2000, n = 5), Small-footed Bat (M. leibii: 2000, n = 1) and Red Bat (Lasiuris borealis: 2000, n = 1). The majority (n = 50; 77%) of bats identified in both years combined were adult, non-scrotal males (Table 1). Eleven nonreproductive adult females were also captured, and the sex of 16 bats was not determined. Since we did not band the bats, it is possible that some individuals were cap- tured repeatedly throughout the study. Marsh (1998) banded 430 bats captured in the summers of 1995 and 1996 at a cave in western Maryland and observed very low recapture success (2.8%). Nonetheless, we were not able to determine if differ- ent nights with a single capture represented the same bat or several different bats over the study period. The maximum number of Big Brown Bats, Eastern Pipistrelles, and Little Brown Bats captured on a given night was 6, 12, and 2, respectively. Although the number of bats using the mine was small (however, we note that we observed more bats than were actually captured avoiding our net and fly- ing into the mine on most nights), interspecific sea- sonal variation in the use of the night roost was evi- dent in 2000 (Figure 1). No bats were captured on two occasions in spring 2000 (4 April, 5 May), likely because the weather was cold and rainy. The mine was used fairly regularly from early-spring to late- summer by a small population of Big Brown Bats. This was expected because this species forages Capture time (minutes after sunset) Juvenile ? Range Mean + SD 2 6 55-323 173.2250 1 6 124-343 238.2 + 61.4 1 4 70-294 2105 + S14 318 213 nightly along the access road and forest clearing adjacent to the mine. The Eastern Pipistrelle was the second most abundant species captured; it exhibited a distinct seasonal use of the mine. Individuals were caught sporadically until late August when 12 Pipistrelles were captured in one night. Many Pipistrelles (more than were actually captured) were observed throughout the night flying around the entrance of the mine. This is consistent with the “swarming” behavior described by Mumford and Whitaker (1975), and may reflect the return of a resi- dent hibernating population to the area. A small number of Little Brown Bats also used the mine spo- radically. Small numbers (<5) of each of these three species have been recorded hibernating in this mine in previous years (D. Feller, personal communica- tion), and it is possible that individuals captured in our study (especially the later part of the study peri- od) subsequently hibernated in the mine. Although each was represented by a single individ- ual, the presence of the Small-footed Bat and Red Bat at the mine is notable. Little is known about the roost- ing habits of Small-footed Bats in the region (Merritt 1987), and the species is listed as threatened in Pennsylvania (Kirkland and Krim 1990). Small-footed Bats are thought to roost deep in rock crevices (Merritt 1987) and are easily overlooked during cen- sus (D. Feller, personal communication). Numerous drill holes and crevices in the mines at RTM provide adequate hibernacula for Small-footed Bats (S. J. Agosta, personal observation). The Small-footed Bat we captured on 9 September suggests that mines also may be used as night roosts by this elusive species. The capture of a Red Bat in August is also of interest because this tree-roosting species rarely uses caves or mines (Merritt 1987); our study site was well-forested and appeared to provide adequate tree-roosting oppor- tunities. Mumford and Whitaker (1975) captured seven Red Bats in July-September at the entrance of a cave in Indiana. It is unknown why Red Bats occa- sionally use caves and mines, but it appears to occur most often in late summer (Webster et al. 1985). For all species combined, nightly capture success (number of bats captured / trap h) at the mine was not correlated with relative humidity (Spearman rank, r = 0.30, P > 0.05), but was positively correlat- 2002 =a = = = oO NM AF O ” toi fe =) ~ Be q O Le Oo 2) z AGOSTA, KUHN, AND MorRTON: BAT ROOST IN MARYLAND 391 M. lucifugus O P. subflavus @ E. fuscus 24-Mar 15-Apr 03-May 04-Jun 22-Jun 06-Jul 28-Jul 10-Aug 23-Aug 07-Sep DATE FiGuRE |. Seasonal use of an abandoned mine used as a night roost by three species of bats in western Maryland during the 2000 foraging season. ed with ambient temperature (r = 0.70, P < 0.05; Figure 2). Many studies have reported a positive relationship between bat foraging activity and ambi- ent temperature (e.g., Lacki 1984; Hayes 1997; Gaisler et al. 1998). Anthony et al. (1981) found an inverse relationship between ambient temperature and the length of night-roosting bouts by female Little Brown Bats, and a positive relationship between ambient temperature and total insect densi- 2.5 1.5 BATS / TRAP HOUR 0.5 6 8 10 12 14 16 18; * 20 22 24 AMBIENT TEMPERATURE (°C) FIGURE 2. Relationship between capture success (number of bats / trap hour) at the abandoned mine night roost and ambient temperature on 19 nights from 4 April to 7 September 2002. Ambient temperature repre- sents the average of three nightly measurements. ty. These data indicate that foraging activity by Little Brown Bats decreases at low temperatures when foraging conditions are poor. In our study, it is possible that the relationship between capture suc- cess at the night roost and ambient temperature reflected different levels of foraging activity in response to variable insect densities (i.e., many bats remained in day roosts at low temperatures). Our limited data also support the findings of Grinevitch et al. (1995) that male and nonreproductive female Big Brown Bats do not forage at ambient tempera- tures < ~12°C (Figure 2). More work is needed on the night-roosting behav- ior of temperate-zone bats (Adam and Hayes 2000). In many areas, night roosts need to be located and documented for this research to progress. Although the data presented here were collected from a single mine, we emphasize the potential value of aban- doned mines to bats, and efforts to salvage and pro- tect these structures should continue or be initiated (see Pierson 1998). The data presented here suggest that abandoned mines may be important resting- places for foraging bats in western Maryland; how- ever, more data are needed to assess their potential significance. Acknowledgments We thank R. Barry and A. Kutay for critical com- ments on an earlier version of this manuscript. We are also grateful to M. Dzialak, A. Moyer, D. Feller, A. Sucke, and K. Kalasz for their help with field- aoe work, and R. Raesly for assisting SJA in locating mine Number 3. Permission to work at Round Top Mountain was provided to SJA by the C&O Canal National Historical Park. Literature Cited Adam, M. D., and J. P. Hayes. 2000. Use of bridges as night roosts by bats in the Oregon Coast Range. Journal of Mammalogy 81: 402-407. Agosta, S. J. 2001. Feeding ecology of the big brown bat (Eptesicus fuscus) in Pennsylvania and western Mary- land. M.S. thesis, Frostburg State University, Frostburg, Maryland. Anthony, E. L. P., M. H. Stack, and T. H. Kunz. 1981. Night roosting and the nocturnal time budget of the little brown bat, Myotis lucifugus: effects of reproductive sta- tus, prey density, and environmental conditions. Oecologia 51: 151-156. Barclay, R. M. R. 1982. Night roosting behaviour of the little brown bat, Myotis lucifugus. Journal of Mammalogy 63: 464-474. Gaisler, J., J. Zukal, Z. Rehak, and M. Homolka. 1998. Habitat preference and flight activity of bats. in a city. Journal of Zoology, London 244: 439-445. Gates, J. E., G. A. Feldhamer, L. A. Griffith, and R. L. Raesly. 1984. Status of cave-dwelling bats in Mary- land: importance of marginal habitats. Wildlife Society Bulletin 12: 162-169. Grinevitch, L., S. L. Holroyd, and R. M. R. Barclay. 1995. Sex differences in the use of daily torpor and for- aging time by big brown bats (Eptesicus fuscus) during the reproductive season. Journal of Zoology, London 235: 301-309. Hayes, J. P. 1997. Temporal variation in activity of bats and the design of echolocation-monitoring studies. Journal of Mammalogy 78: 514-524. Hirshfeld, J. R., Z. C. Nelson, and W. G. Bradley. 1977. Night roosting behavior in four species of desert bats. Southwestern Naturalist 22: 427-433. Kirkland, G. L., and P. M. Krim. 1990. Survey of the statuses of the mammals of Pennsylvania. Journal of the Pennsylvania Academy of Science 64: 33-45. Kunz, T. H. 1982. Roosting ecology. Pages 1-55 in THE CANADIAN FIELD-NATURALIST Vol. 116 Ecology of bats. Edited by T. H. Kunz. Plenum Press, New York. Lacki, M. J. 1984. Temperature and humidity-induced shifts in the flight activity of little brown bats. Ohio Journal of Science 84: 264-266. Lewis, S. E. 1994. Night roosting ecology of pallid bats (Antrozous pallidus) in Oregon. American Midland Naturalist 132: 219-226. Marsh, B.D. 1998. Prehibernal weight gain and activity of temperate cave-dwelling bats in western Maryland and southwestern Pennsylvania. M. S. thesis, Frostburg State University, Frostburg, Maryland. | Merritt, J. F. 1987. Guide to the mammals of Pennsyl- vania. University of Pittsburgh Press, Pittsburgh. Mumford, E., and J. O. Whitaker, Jr. 1975. Seasonal activity of bats at an Indiana cave. Proceedings of the Indiana Academy of Science 84: 500-507. O’Shea, T. J., and T. A. Vaughan. 1977. Nocturnal and seasonal activities of the pallid bat, Antrozous pallidus. Journal of Mammalogy 58: 269-284. Perlmeter, S. I. 1996. Bats and bridges: patterns of night roost activity in the Willamette National Forest. Pages 132-150 in Bats and forests symposium, October 19-21, 1995, Victoria, British Columbia. Edited by R. M. R. Barclay and R. M. Brigham. Research Branch, Ministry of Forests, Victoria. Pierson, E. D. 1998. Tall trees, deep holes, and scarred landscapes: conservation biology of North American bats. Pages 309-325 in Bat biology and conservation. Edited by T. H. Kunz and P. A. Racey. Smithsonian Institution Press, Washington D.C. Pierson, E. D., W. E. Rainey, and R. M. Miller. 1996. Night roost sampling: a window on the forest bat commu- nity in northern California. Pages 151-163 in Bats and forests symposium, October 19-21, 1995, Victoria, British Columbia. Edited by R. M. R. Barclay and R. M. Brigham. Research Branch, Ministry of Forests, Victoria. Webster, W. D., J. F. Parnell, and W. C. Biggs, Jr. 1985. Mammals of the Carolinas, Virginia, and Maryland. University of North Carolina Press, Chapel Hill. Received 17 January 2001 Accepted 22 May 2002 Relative Abundance, Habitat Use, and Breeding Status of Birds in Aulavik National Park, Banks Island, Northwest Territories J. DAviIpD HENRY! and MICHELLE MIco2 !Parks Canada, P.O. Box 5495, Haines Junction, Yukon Territory YOB 1L0 Canada 2Western Canada Service Centre, Parks Canada, 145 McDermot Avenue, Winnipeg, Manitoba R3B OR9 Canada Henry, J. David, and Michelle Mico. 2002. Relative abundance, habitat use, and breeding status of the birds in Aulavik National Park, Banks Island, Northwest Territories. Canadian Field-Naturalist 116(3): 393-407. During the summers of 1995 and 1996, we inventoried bird populations of Aulavik National Park (73°N, 119°W), located on northern Banks Island in Canada’s Arctic Archipelago. Our study added four new species, bringing the total of bird species to 44 for Aulavik and 74 for Banks Island as a whole. We also documented 21 new breeding records in the park, and the known breeding distribution of Semipalmated Plover (Calidris pusilla), Purple Sandpiper (Calidris maritima) and Buff-breasted Sandpiper (Tryngites subruficollis) were extended. Point-counts were carried out in three different study areas of the park and used to determine relative abundance for 13 species. Lapland Longspur (Calcarius lapponicus) was the most common species. Lapland Longspurs, Horned Larks (Eremophilia alpestris), Glaucous Gulls (Larus hyper- boreus), and Black-bellied Plovers (Pluvialis squatarola) were significantly associated with a few different habitat types. No habitat type had notably more bird species as compared to the other types, with the exception of sparsely vegetated ground which supported fewer species. Field observations, lower average wind speed in hummocky tundra and the pres- ence of a high number of owl castings and mammal scat in deep, wind-protected coulees suggested that both small and large birds as well as certain mammals seek out sheltered microhabitats during high winds. It is suggested that park opera- tions should not interfere with the access that birds and other wildlife have to these potentially important microhabitats. Key Words: birds, breeding distribution, abundance, habitat associations, microhabitats, wind shelter, Banks Island, Aulavik National Park, Arctic, Northwest Territories During the summers of 1995 and 1996, a team of Parks Canada scientists and Inuvialuit inventoried the biological and archaeological resources of Aulavik National Park, a park established in 1992. This report describes the occurrence, breeding status, relative abundance, and habitat use patterns of the birds of the park. Study Area Aulavik National Park is a large, remote, protect- ed area in the northern portion of Banks Island in Canada’s Arctic Islands (see Figure 1). Aulavik encompasses 12 275 square kilometers of the Western Arctic Lowland Natural Region (Parks Canada 1990). Banks Island Migratory Bird Sanctuary Number 2 is within the park (see Figure 1) and is managed jointly by the Canadian Wildlife Service and Parks Canada. Aulavik National Park contains a diversity of landscapes — upland plateaus, unvegetated polar desert areas, wetlands with frost polygon features, sheer cliffs along the coast and southeast of Mercy Bay, and the lush val- ley of the Thomsen River (Gray and Alt 1997*). Most of Banks Island has probably been free of glaciation for millions of years or never glaciated (Zoltai et al. 1980*). All land surfaces of the park are underlain by permafrost. The average thickness *See Documents Cited section wy) of the active layer during August is approximately 50 cm, shallower on peaty soils and deeper on well- drained soils (Gray and Alt 1997*). At least 169 species of vascular plants have been recorded in Aulavik National Park (Raillard in press). The vege- tation of the park is frequently characterized by polar semi-desert and desert community types (Bliss et al. 1984; Bliss and Svoboda 1984). Moister sites sup- port more productive graminoid shrub tundra and wet sedge meadows. Ferguson (1991) described eight habitat types in the Muskox River valley. Dur- ing our study these habitat types were verified in the Muskox, Eames, and Thomsen river valleys and were used to classify habitat for this project. Methods Field work for this bird inventory was carried out from 26 June to 2 August 1995 and 8 to 18 July 1996. In order to compare weather from these two periods to long-term averages, basic weather data were collected twice each field day (approximately 07:00 h and 19:00 h) including total precipitation, ambient temperature, wind speed, and wind direc- tion. Wind speed was recorded with a hand-held anemometer (Dwyer Wind Meter, Dwyer Instru- ments Inc., Michigan City, Indiana). Cloud cover was described using standard aviation terminology. Cloud, temperature, and wind conditions were also described for each point-count. 394 Ten different areas of the park were inventoried for birds (see Figure 1). Every field day a list of bird species with approximate numbers was compiled. One to 12 daily checklists were compiled for each area. Number of checklists depended on how many days observers spent in each area which was deter- mined by logistic support and other research occur- ring there. More time was spent in the more produc- tive areas, and birds from these areas were better represented on the checklists than birds from less productive areas. Breeding records were recorded only for sightings of an active nest with eggs or off- spring, or of an adult bird with flightless young. From these daily checklists, bird species were grouped in three relative abundance classes: common (species recorded on 50 percent or more of check- lists), uncommon (species recorded on 49 to 20 per- cent of checklists), or rare (species recorded on less than 20 percent of checklists). Because several vari- ables influence the frequency with which a bird is recorded (e.g., conspicuousness, distance its ‘song can be heard, reaction to humans), relative frequency of bird species, as determined by daily checklists, cannot be defined more precisely (Scotter et al. 1985). Point-counts were carried out on three biologically productive study areas within the park: Eames River, Muskox River, and the central portion of the Thomsen River (see Figure 1). These are referred to as study areas. A standard point-count methodology for open environments (100 m radius, 5 minute observation time) was employed during this study (Ralph et al. 1995). Point-counts were carried out between 05:00 and 13:00 h, but not during inclement weather (heavy wind, fog, snow or rain). Point- counts were randomly located within a single habitat type, and perimeters of point-counts were separated from each other by at least 200 m. The centre of the point-count was marked, as well as two opposite points on the perimeter, each 100 m from the centre. The observer returned to the centre, waited approxi- mately | minute to reduce disturbance effects, and then for the next 5 minutes mapped all birds seen or heard on the 100 m radius circular area as well as birds seen or heard outside the plot or flying over the plot (see Henry and Mico 1997*, for details). Move- ments and interactions of birds present on the plot were recorded and mapped in order to reduce the chance of recording the same bird more than once. Observers made special efforts not to record on more than one count birds that could be seen or heard over long distances (e.g., Sandhill Cranes, Common Ravens) [see Appendix for scientific names]. A total of 20 to 35 point-counts was recorded for each of eight habitat types (Ferguson 1991) on each study: area. In total, 764 point-counts were recorded. During the two field seasons, none of the point- counts were revisited. THE CANADIAN FIELD-NATURALIST Vol. 116 ras The point-count data were analyzed to examine relative abundance and habitat associations. Only 13 species were recorded on at least five point-counts and on at least two study areas. These species were judged to be recorded frequently enough to analyze their abundance and habitat associations. The rela- tive abundance and habitat association data were not normally distributed; thus Friedman’s Nonpara- metric Two-Way ANOVA test was used to analyze the data (a = 0.05). If the null hypothesis that on the three study areas bird species were equally abundant (or the null hypothesis that bird species were equally abundant on all habitat types) was rejected, a multi- ple comparison procedure (modified Newman-Keuls test; Daniel 1990) was utilized (setting experiment- wide error at a = 0.10; Daniel 1990). This latter test was used to determine which species exhibited sig- nificantly different abundance values across the three study areas and across the habitat types. To assess the extent that hummocky tundra can provide shelter for small birds during windstorms, wind speed was measured with a hand-held ane- mometer on days that exhibited an average wind speed of 15 k/h at 1.5 m above ground. Wind speed measurements at 35 different locations were taken at several different heights: 150 cm above ground, 100 cm, 50 cm, 20 cm, 10 cm, 0 cm as well as in the bottom of microdepressions between hummocks. Average wind speed measurements were normally distributed and were tested for significant differences using a single factor ANOVA analysis (Sokal and Rohlf 1969). Pellet-count transects tested whether coulees were selectively used by large birds and large mammals as compared to areas of surrounding tundra. Randomly selected paired 100 m transects were set up, one transect in the bottom of a sheltered coulee and the other on nearby flat, open tundra. On each transect, all scats and owl castings within | m of the transect centreline were recorded. Five pairs of transects were sampled for each of three different sheltered coulees. A t-test (n = 15) was used to analyze for sig- nificant differences in scats and castings on the two types of transects (Sokal and Rohlf 1969). Results and Discussion Weather Patterns Research from the nearest permanent weather sta- tions, at Sachs Harbour on Banks Island and Mould Bay on Prince Patrick Island, provided by Mete- orological Services of Canada, Environment Canada (1996*) were used to calculate long-term weather averages (Table 1). Both weather stations are located near the coast whereas the study areas were 15 to 40 km inland from M’Clure Strait. The difference in weather of coastal regions compared to more inland parts of Banks Island has not been studied in detail. Coastal regions may be cooler, foggier, and wetter than more inland locations (Zoltai et al. 1980*). 2002 * ! | | Nangmagvik, j Lake ; ! | t l en eee s | ae oeilien | eS . eS Se * HENRY AND Mico: BIRDS IN AULAVIK NATIONAL PARK 395 \ \ o)%,, A Rive, 1 I . ond Audavik National Park @; Banks [sland ig “4545 oe FiGuRE 1. The location of Banks Island and Aulavik National Park. The centre of the park is located at approxi- mately 73°42’N, 119°20’W. The hatched area on the map is Banks Island Bird Sanctuary Number 2 managed jointly by the Canadian Wildlife Service and Parks Canada. The @ indicates the ten areas of Aulavik National Park where daily checklists of bird species were compiled. The numbers indicate the three study areas of the park where point-counts were carried out. These study areas are as follows: (1) Eames River, (2) Muskox River and (3) central portion of the Thomsen River. The 1995 field season was generally warmer, wet- ter, and calmer and the 1996 field season was warmer and calmer than the long-term averages (Table 1). Thus most of the field work for this project was car- ried out under average to good weather conditions. Two periods (22 July to 2 August 1995 and 18 to 26 July 1996) were characterized by precipitation and/or strong winds. No point counts were carried out dur- ing these periods of inclement weather. Classification of Habitat Types All habitats sampled during this study were classi- fied according to Ferguson (1991) with slight modi- fications relevant to avian species (for details, see Henry and Mico 1997). As listed below, the terrestri- al habitat types progress from wet, low sites with higher aboveground primary productivity to xeric, less productive sites at higher elevations. The habitat types used in this study are: 396 Water Bodies (WB). This habitat type includes open water of lakes, rivers, streams, and tundra ponds. Wet Sedge Meadows (WSM). This type occurs on level, hydric lowlands. It is usually covered by shallow (<10 cm) water and a nearly continuous cover of sedges, especially Carex aquatilis var. stans, and other hydrophytic species. Graminoid Tundra (GRT) occupies mesic to hygric sites in lowlands and on gentle slopes, with continuous cover of graminoid species, and sometimes dwarf shrubs (usually Salix species). Plant growth is most luxuriant on gentle slopes downslope from snowbeds. Graminoid/Dwarf Shrub Tundra (GST). This type is intermediate between graminoid tundra and dwarf shrub tundra in terms of moisture and vegetation. Vegetation cover, characteristically between 75 to 100 percent, is a mosaic with graminoids and mosses dominate in moist depressions and herbs and dwarf shrubs dominate on the drier hummocks. Dwarf Shrub Tundra-(DST) occurs on moist, well- drained, middle and upper slopes as well as solifiuction lobes and terraces. Vegetation cover, characteristically 50 to 75 percent, is dominated by dwarf shrubs, especially Salix arctica and Cassiope tetragona. Small, non-sorted polygons as well as cryoturbated surfaces are often present. Hummocky Tundra (HT). This type is non-sorted earth hummocks covering small to extensive areas. Narrow fur- rows and cracks separating the hummocks support mosses, lichens, herbs, and sometimes dwarf shrubs. The tops and sides of hummocks are dry and often free of vegetation. Dwarf Shrub/Lichen Barrens (DLB). This variable type occurs at all elevations on windblown sites. Typically sites are rapidly drained, consisting of silty to stoney soils. Vegetation, characteristically 25 to 50 percent cover, is dominated by mat and cushion plants. Sparsely Vegetated Ground (SVG) is characterized by less than 10 percent vascular plant cover. This type occurs on cutbanks, mudflats, sand and gravel bars, uplands, wind- blown sites, sand dunes, frost-shattered bedrock, debris- mantled slopes, and bare sandstone. Lists of Bird Species for Aulavik National Park and Banks Island On the basis of literature review and fieldwork, Zoltai et al. (1980*) listed 40 bird species in Aulavik National Park. During eight weeks of fieldwork for this project, we added only four new bird species to the park’s list: Tundra Swan, Red-necked Phalarope, Western Sandpiper, and Purple Sandpiper. Henry and Mico (1997*) provide details of these sightings. THE CANADIAN FIELD-NATURALIST Vol. 116 Zoltai et al. (1980*) reported Gyrfalcons and Thayer’s Gulls in the park area. Wilkinson et al. (1977*) reported Lesser Yellowlegs in the park area, but Zoltai et al. (1980*) considered this species as hypothetical. These three bird species were not observed during this study. Thus in total, 44 bird species have been recorded in Aulavik National Park, 41 of which were observed during this study (Table 2). Zoltai et al. (1980*) summarized all prior sight- ings of bird species for the park as well as for Banks Island. This list was updated (see Appendix), and it now incorporates all documented and reported sight- ings and breeding records up through 2001. Several interesting patterns are evident. Amundsen Gulf, which separates Banks Island from the mainland, is confirmed to be an effective geographic barrier to migrating birds. For example, 143 bird species have been recorded in Ivvavik National Park in the north- ern Yukon Territory (Holcroft Weerstra 1997*) and 127 species in Vuntut National Park (Gray and Alt 2000*; Henry et al. 2002) as compared to 74 bird species for Banks Island and 44 species in Aulavik National Park (see Appendix). The fact that only 44 out of 74 bird species observed on Banks Island have been observed in the park may reflect the limited research effort in the park. It might also reflect the reduced biological pro- ductivity and wider distribution of polar desert habi- tat in the park as compared to other parts of Banks Island (Gray and Alt 1997*; Raillard in press). There are documented sightings for 23 bird species (plus seven species recorded as hypothetical) in other parts of Banks Island that have not yet been observed in Aulavik National Park (see Appendix). This difference includes a variety of species, for example, waterfowl (Ross’ Goose, Northern Pintail, Common Eider, and Red-breasted Merganser); a rap- tor (Short-eared Owl); a shorebird (Whimbrel) as well as a passerine species (Barn Swallow). The Appendix includes single sight records for the Western Sandpiper in Aulavik National Park as well as 13 additional bird species on Banks Island. Because a specimen was collected, a photograph was obtained, or a detailed description was provided, TABLE |. Weather parameters for July 1995 and July 1996 compared to long-term averages for July from the nearest permanent weather stations. Aulavik Study Area Sachs Harbour Mould Bay 1995 1996 1955-1990 July 1948-1990 July 26 June—02 August 19-28 July Long-term Mean Long-term Mean Mean Daily Max. Temp (°C) 12.0 12.8 10.0 6.5 Mean Daily Min. Temp (°C) 2.1 4.0 3.0 Re | Average Wind Speed 13.0 10.0 n/a 18 Total Precipitation (mm) 125.0 20.0 bn 14.2 2002 HENRY AND Mico: BIRDS IN AULAVIK NATIONAL PARK 397 TABLE 2. Relative abundance, based on daily checklists, and breeding status of 41 avian species observed in Aulavik National Park during the summers of 1995 and 1996. Common species were recorded on 50 percent or more of the daily checklists, uncommon on 20 to 49 percent, and rare on less than 20 percent of the checklists. Breeding records listed as new were established as a result of field observations made during this study. Species Relative Abundance Breeding Record Red-throated Loon* Rare Pacific Loon* Uncommon X new Yellow-billed Loon* Rare X new Snow Goose* Rare Canada Goose Rare X new Brant* Uncommon X new Tundra Swan Rare King Eider** Uncommon X new Long-tailed Duck* *Rare X new Rough-legged Hawk Uncommon Xx Peregrine Falcon** Rare x Willow Ptarmigan Rare X new Rock Ptarmigan** Rare x Sandhill Crane Common X new Black-bellied Plover** Common X new American Golden-Plover** Uncommon X new Semipalmated Plover Rare Ruddy Turnstone** Rare X new Purple Sandpiper* Rare X new Sanderling* Rare os Semipalmated Sandpiper Uncommon x Western Sandpiper Rare Least Sandpiper Rare White-rumped Sandpiper** Rare X new Baird’s Sandpiper** Uncommon xX Pectoral Sandpiper* Uncommon X new Buff-breasted Sandpiper Rare X new Red-necked Phalarope Rare Red Phalarope** Rare X new Pomarine Jaeger* Rare Parasitic Jaeger* Rare Long-tailed Jaeger* Common X new Glaucous Gull** Common xX Sabine’s Gull* Rare X Arctic Tern** Uncommon X new Snowy Owl* Uncommon Xx Common Raven* Uncommon X new Horned Lark* Common Xx American Pipit* Rare xX Lapland Longspur ** Common X new Snow Bunting** Uncommon X new *Species is known to occur in both Truelove Lowland, Devon Island (Pattie 1990) and Aulavik National Park (this study). **Species is known to occur and breed regularly in both Truelove Lowland, Devon Island (Pattie 1990) and Aulavik National Park (this study). these 14 species records are considered to be accurate, and the Appendix lists these species as accidental occurrences on Banks Island. These species include: Northern Fulmar, Spectacled Eider, Western Sandpiper, Black Guillemot, Tree Swallow, Barn Swallow, Black-billed Magpie, Northern Waterthrush, American Redstart, Common Redpoll, American Tree Sparrow, Harris’s Sparrow, White- crowned Sparrow, and Fox Sparrow. The Appendix lists another seven bird species as hypothetical occurrences on Banks Island, and these species need confirmation. These species include: Greater White- fronted Goose, American Wigeon, Lesser Yellow- legs, Dunlin, Black-legged Kittiwake, Thick-billed Murre, and Dovekie. Large rocky and sandy barren areas, often described as polar deserts, are widely distributed on several Canadian High Arctic Islands (Bliss et al. 1984) and occur often in Aulavik National Park. These areas are at least a square kilometer in size, sparsely vegetated and include windblown ridges, high rocky uplands, sand dunes, talus slopes below 398 cliffs, as well as large mud flats and gravel bars in or adjacent to rivers. In Aulavik they cover 11.7 percent of the total area (Raillard in press). Birds are infre- quently observed in these polar desert areas. For example, during mid-July 1996, six hours of survey- ing such areas west of Nangmagvik Lake (see Figure 1) yielded only four birds — one Long-tailed Jaeger, one Horned Lark, and two Lapland Longspur. Sightings of approximately one bird per hour were typical for the driest of these polar desert areas. Bliss and Svoboda (1984) described the wide dis- tribution of polar semi-desert habitat among Canadian Arctic Islands north of 70° latitude. In Aulavik National Park 24.5 percent of the park area is classified as polar semi-desert (Raillard in press). The ground surface of these areas is stony or covered with sand and clay, and plant cover is between 10 and 50 percent (see Raillard in press, for details). Together, polar desert and polar semi-desert areas constitute 36.2 percent of Aulavik National Park. These areas are characterized by few bird species and low avian densities. Checklists of Birds Observed on Each Field Day Observers compiled separate checklists of birds each day in the field. Six out of 41 species (15 per- cent) were commonly observed; 12 species (29 per- cent) were uncommonly observed, and 23 species (56 percent) were rarely observed in these areas of the park (Table 2). Patterns observed during the two summers were relatively consistent. Thirty-eight bird species were observed in the 1995 field season, and 35 species in the 1996 field season. The Red-throated Loon, Tundra Swan, Sabine’s Gull, Sanderling, Western Sandpiper, and American Pipit were record- ed only in 1995. The Semipalmated Plover, Red- necked Phalarope, and Least Sandpiper were record- ed only in 1996. Checklists were compiled for ten areas in the park; thus, many other areas still need to be studied. However, as eight weeks of fieldwork found only four new bird species, these results suggest that the avifauna of Aulavik National Park is fairly well doc- umented. The 44 species recorded in the park seem to be typical bird species observed in the Middle Arctic region (Pielou 1994). During 16 years of censusing breeding bird populations on Truelove Lowland, Devon Island, Nunavut (75°41°N, 84°35’W), Pattie (1990) recorded 43 species of birds, 18 as regular breeders. The avian communities from these two locations are similar. Seventy percent (31 out of 44 species) occurs in both Aulavik National Park and Truelove Lowland (these species are marked as * or ** in Table 2, plus Thayer’s Gull, see Appendix). Of the 18 regular breeders in Truelove Lowland, 14° species (78 percent) breed in Aulavik National Park. These similarities suggest that the same avian species occur over much of the Middle Arctic and THE CANADIAN FIELD-NATURALIST Vol. 116 breed there. Other localities in the Middle Arctic should be examined for this assemblage of species. Breeding Bird Records for Aulavik National Park As a result of their literature review and field- work, Zoltai et al. (1980*) listed 12 breeding bird species in the park area (see Appendix). Using more exact criteria for breeding records (see Methods), our study documented breeding by 21 additional species in Aulavik National Park (Table 2). For three species, this study also extends the known detailed breeding range as presented by Godfrey (1986) and updated by Sibley (2000). These three species are: Semipalmated Sandpiper, Purple Sandpiper, and Buff-breasted Sandpiper. In summary, 33 of the 44 bird species (75 percent) known to occur in Aulavik National Park are now known to breed there (Table 2, plus Thayer’s Gull in the Appendix). Nests of Peregrine Falcons, arctic race, containing flightless offspring were observed on cliffs adjacent to a river and a stream within the park. Gyrfalcon Bluff is the only location on Banks Island where Gyrfalcons have been documented to nest and raise young. However, Gyrfalcons have not been observed nesting there since 1979. A review of sightings of Gyrfalcons (Henry and Mico 1997*) supports the view that Gyrfalcons are rare in Aulavik National Park and may not nest there at present. Cliff nesting sites for Peregrines, Gyrfalcons and Rough-legged Hawks occur infrequently over much of the interior of Aulavik National Park and may be a limiting factor for these populations. For example, on three occasions Rough-legged Hawks nested and laid eggs on flat- topped boulders that were 2 m or less above ground level. These nests seemed easily accessible by Arctic Foxes and other predators. If future field research sup- ports that cliff nesting sites for raptors are rare in the park, these sites may warrant special management action to protect them from human disturbance. Relative Abundance of Common Bird Species Daily checklists give a coarse indication of relative abundance of bird species (Table 2). The point-count method (Ralph et al. 1993, 1995) is a more quantita- tive technique, and it was used to estimate the relative abundance and examine the habitat association pat- terns for common bird species. Point-counts were used on three study areas (see Figure 1): Eames River area (centered at 74°06.2148N, 120°45.07°78W), Muskox River area (centered at 73°45.6778N, 120°34.849W), and the central portion of the Thomsen River area (centered at 73°13.8498N, 119°32.4228W). There are significant differences among the relative abundances of 13 bird species (Friedman’s ANOVA, P < 0.05). The Lapland Longspur was the most abundant bird species on these three study areas. Its relative abundance was at least three to five times higher than the next most numerous bird species, Black- bellied Plover. 2002 HENRY AND MIco: BIRDS IN AULAVIK NATIONAL PARK 399 Black-bellied Plover was consistently ranked and Brant) or even rare (Red Phalarope) as deter- between second and fourth in all estimates and aver- mined by daily checklists (Table 2). The point-counts aged the second most common bird species on the __ were carried out in biologically productive areas of study areas. The relative abundances of Horned Lark the park whereas the daily checklists were from areas and Baird’s Sandpiper are significantly lower varying from polar deserts to productive river valleys. (Newman-Keuls test, P < 0.10). Their relative abun- dances are nearly equal, and they tied as the third most common species on the study areas. Relative abundance of Horned Lark is more variable than that of Baird’s Sandpiper. Long-tailed Jaeger was fourth in abundance but shows variation, being more abundant on the Muskox River and Thomsen River study areas than on the Eames River study area. The relative abundance of Pectoral Sandpiper and Sandhill Crane is significantly lower than that of Long-tailed Jaeger, and these two species are tied for fifth in abundance on the study areas. Red Phalarope and Glaucous Gull are tied for S©4d8* meadow (WSM), however, had only interme- sixth in abundance, and they are only slightly less diate avian abundance. Hummocky tundra (HT) is abundant than Pectoral Sandpiper and Sandhill Crane. ™€SIC and has an intermediate avian density. Finally, American Golden-Plover, King Eider, and Pacific Sparsely vegetated ground (SVG), dwarf shrub- Loon are tied for seventh in abundance. The _ lichen barrens (DLB), and dwarf shrub tundra (DST) American Golden-Plover and Pacific Loon show lit- ate dry habitats with low density of birds. tle variation in abundance among study areas. The Vegetation cover (which is also related to available King Eiders were mostly females raising young on surface moisture) also appeared to be a good predic- the inland areas of Banks Island while the males dur- tor of density of birds in certain habitats: terrestrial ing summer tend to congregate along the coast. sites with 75 percent or less vegetation cover had the The Brant is the least common of these 13 avian lowest avian densities. In descending order of vegeta- species. However, it was relatively common on the _ tion cover and avian abundance, these sparsely vege- Muskox River study area, where a flock of overahun- _ tated habitat types are ranked DST > DLB > SVG. dred birds was observed repeatedly on a small lake. As discussed above, bird densities respond to sur- The composition and relative abundance of the face moisture and habitat productivity: with some avian community vary as habitats and ecological con- exceptions, habitats with high surface moisture ditions in different parts of Aulavik National Park exhibit higher bird densities. Point-counts were also change. Some of the relatively common bird species, utilized to determine habitat association patterns as determined by point-counts, were found to be _ based on Ferguson’s (1991) habitat types for each of uncommon (Baird’s Sandpiper, Pectoral Sandpiper, the 13 common bird species. Nine out of 13 species American Golden-Plover, King Eider, Pacific Loon showed no significant preference for any of these Habitat Association Patterns For each study area, overall avian density (mean number of individual birds of all species per point count) was calculated for each habitat type. Certain habitat types supported significantly higher density of birds than others (Friedman’s ANOVA, P < 0.05). The data showed a relationship between available surface moisture and avian density: wet habitats had the highest abundance of birds. Water bodies (WB), graminoid-dwarf shrub tundra (GST), and graminoid tundra (GRT) had the highest density of birds. Wet TABLE 3. List of mean + standard deviation of birds from 764 point-counts with 100 m radii. Densities given for the 13 most common bird species in Aulavik National Park according to the eight habitat types (n = number of individual birds observed). Bird species names are abbreviated by using first three letters of separate words in the AOU name for the bird (e.g., BLA PLO means Black-bellied Plover). WB WSM GRT GST DST HT DLB SVG Species (n=68) (n=91) (n=100) (n=101) (n=85) (n=104) (n=96) (n=120) oo PAC LOO 2+ 42 06+.09 . 032.06 .03+.03 0.00. 02 + .03 .0.00 03+.06 0.349 BRA 51+ 9.6 D210. i233 024.04 034.07 0.00 0.00 .0.00 0.371 KIN EID 98+ 1.1 1624.20 (192.39 §42.11 0.00 0.00 28 + 34 0.00 0.234 SAN CRA .07 + .10 eae «tet OS.) 24.33 . O05 2°10 07+.11 .16+.29 .09+.06 0.826 BLA PLO ‘26 35 31.14 28+ 24 47+A9 292.19 134.09 122.08 .06+.07 0.085 AME GOL 01 +02 02203 252 07 32% AZ 0.00 04+.03 05+.07 O12+.02 0.738 RED PHA .07 + .11 2624.27 03 £107" .01 = .03 0.00 0.00 0.00 0.00 0.397 BAI SAN .O5 + .03 164 (10) 21% 22 06% 075° 17 £22 182.16 .12+.14 .08+.05 0.467 PEC SAN 12+ .14 05+ 07 .08+.16 .01+.03 0.00 0.00 0.00 0.00 0.787 LON JAE 20 +416 A418) 182513 302.24 272.21 164.19 432.42 .142.11 0.304 GLA GUL 30 + 37 102.10 .24+.19 24.23), ..15+.27 07 +.11 09+ .04 02+.02 0.064 HOR LAR 0.00 O1+,01.. .05.2.08.. .17+.06.. .12+.08 172.12 .172.09 .10+.13 0.062 LAP LON 14+ 89 LO ie kee Dal SR a Hi ES p Nie SE RE I I oe I 9) 400 habitat types (Table 3, Newman-Keuls test, P>0.10). At this level of habitat classification, these species appear to be habitat generalists. Lapland Longspur, Horned Lark, Glaucous Gull, and Black- bellied Plover were significantly associated with cer- tain habitat types (Table 3, Newman-Keuls test, P < 0.10) as follows: Lapland Longspurs were most abundant on HT and GST but seemed least abundant on or immedi- ately next to WB, SVG, and DLB. Horned Larks were similar to but not identical with Lapland Longspurs. Horned Larks were most abundant on HT, GST, and DLB. However, this species seemed least abundant on or next to WB or on WSM and GRT — all habitat types with high surface moisture. Glaucous Gulls are a wide-ranging species, espe- cially while foraging. However, they were most commonly observed on or next to WB where they nested on small islands in medium to large tundra ponds. They were also frequently observed in mesic sites such as GRT and GST, but they were rarely observed on SVG. Black-bellied Plovers were rarely observed on SVG. This species appeared to be adapted to a wide range of moisture conditions, being most commonly observed quite near WB or on WSM, GRT, GST, and DST. During his sixteen-year study of the birds of Truelove Lowland, Pattie (1990) documented a neg- ative correlation between the decreasing population of Black-bellied Plovers and increasing American Golden-Plovers. On several occasions he observed American Golden-Plovers pursuing Black-bellied Plovers and suggested that there may be competition between these species. On the Aulavik study areas, Black-bellied Plovers were four times more abun- dant than American Golden-Plovers, and no agonis- tic interactions between these species were observed. Although American Golden-Plovers showed no sig- nificant habitat associations, they were most fre- quently observed on GRT, whereas Black-bellied Plovers were observed on a wide range of moist habitat types. It may be that competitive interactions between these species do not occur until the popula- tion of American Golden-Plovers reaches a certain threshold density. The number of bird species observed in each habi- tat type was examined. Seven of the habitats showed no significant differences in the number of bird species observed (Newman-Keuls test, P > 0.10). SVG supported fewer bird species (Newman-Keuls test, P < 0.10). SVG on river banks and gravel bars of the central portion of the Thomsen River support- ed more bird species than similar habitat on the Eames River or Muskox River. The Thomsen River is one of the largest rivers in the park and may have less permafrost and more invertebrates associated with its gravel bars and river banks than other rivers and streams in the park. THE CANADIAN FIELD-NATURALIST Vol. 116 rs Shelter During Storms Finding shelter from the wind can be critically important to many plants and animals on the tundra. During a typical summer in the Middle and High Arctic, strong windstorms accompanied by near- freezing temperatures and rain or snow can last a week or longer before abating (Pielou 1994). Reducing heat loss and minimizing energy expendi- ture during storms may be critical to survival of small birds and their offspring (Rodrigues 1994). Wiersma and Piersma (1994) and Rodrigues (1994) discussed the importance of energy-conserving behaviours by birds, such as the selection of nest sites. Hummocky tundra provides shelter for the nests of many arctic- breeding passerines (Ehrlich et al. 1988). During this study several pairs of Lapland Longspurs and Snow Buntings were observed to nest in hummocky tundra or other small microdepressions. Terrain such as hum- mocky tundra may also be critical in providing shelter for all adult passerines during wind and rainstorms. Aboriginal people and naturalists have often observed that small birds on the tundra seem to take shelter dur- ing periods of high winds (e.g., Rand 1948; Porsild 1951). When a person walks through hummocky ter- rain during such times, Lapland Longspurs, Horned Larks and other small birds often explode into flight out of small depressions in the tundra. To test the potential shelter provided by these microhabitats, wind profiles were measured at 35 locations when the average wind speed was approxi- mately 15 k/h (see Methods and Figure 2). Average wind speed at the bottom of microdepressions was 1/20th of what it is at 150 cm above ground and near- ly 1/10th of wind speed at ground level (single factor ANOVA, P < 0.0001). Protection from wind is not simply a function of depth but also a function of verti- calness of walls and overhang of uppermost edges of microdepression. Nine microdepressions with these favourable features were measured, and wind speed was reduced to approximately 1/1000th of what it was 150 cm above ground. Consequently, by carefully choosing the microdepression where they nest or those they occupy during a windstorm, small passer- ines might reduce heat loss considerably. Larger birds (e.g., cranes, geese, and ptarmigans) and mammals that do not den underground (e.g., muskoxen and caribou) also might seek shelter during periods of high wind and precipitation. In Aulavik National Park, deep and narrow coulees occasionally form along small creeks near their confluence with larger rivers. During a twelve-day period of wind, rain, and snow in early August 1995, JDH observed a pair of Sandhill Cranes, a Common Raven, and a ptarmigan standing in the bottom of one of these ’ coulees for more than five hours. These birds were feeding only infrequently. Most of the time they sim- ply stood quietly, within 10 m of each other, apparent- ly seeking shelter from the storm. 2002 r— = = — 3 vo oe o. a z = 150 100 50 centimeters above ground HENRY AND Mico: BIRDS IN AULAVIK NATIONAL PARK 401 20 10 Ground Bottom of level . ‘ microdepression FIGURE 2. Wind speed (mean and standard error) at various distances above the ground and in the bottom of microdepressions of hummocky tundra. Paired pellet-count transects indicate that large birds and certain mammals defecate more in these coulees than in open tundra areas immediately adja- cent to them. This distribution of scat suggests that these animals spend selectively more time there. In 15 different locations, droppings of large birds and mammals were from five to ten times more numer- ous in sheltered coulees than in adjacent tundra areas (K.outees = 22:1 scats; x, 4, = 1.8 scats; t = 6.346, < 0.001). Wind speed measurements suggested that narrow, steep-walled coulees oriented perpen- dicular to prevailing winds offer considerable shelter during windstorms. Such coulees were extremely rare in the ten areas of the park examined. It is rec- ommended that park staff and visitors avoid using these coulees, and that no park facilities are devel- oped near these areas so as not to interfere with free access by large birds and mammals to these poten- tially important microhabitats. Acknowledgments This research project received support from the Western Canada Service Centre and the Western Arctic Field Unit of Parks Canada. Martin Raillard, Steve Catto, Yvonne Rowland, and Vicki Sahantien were especially helpful, and they contributed valu- able bird observations during the two field seasons. The project received financial support from the Green Plan Ecological Integrity Fund. Documents Cited [marked with an * in the text] Meteorological Services of Canada, Environment Can- ada. 1996. 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Dobkin, and D. Wheye. 1988. The Birder’s Handbook. Simon and Schuster Inc. New York, New York. 785 pages. Ferguson, R.S. 1991. Detection of muskox habitat on Banks Island, Northwest Territories, Canada, using Landsat Thematic Mapper data. Arctic 44: 66-74. Godfrey, W. E. 1986. The Birds of Canada (Revised edi- tion). National Museum of Natural Sciences Bulletin 203: 1-428. Henry, J. D., B. Farkas, and R. Markel. 2002. Bird sight- ings and breeding records from Vuntut National Park and surrounding areas, 1997-2001. Yukon Warbler 8 (1): 6-9. Holyoak, D. T. 1983. Notes on the birds of southwest Banks Island, Northwest Territories, Canada. Bulletin of the British Ornithological Club 103: 37-39. Latour, P., and L. M. Allison. 1976. Four new bird records for Banks Island, Northwest Territories. Can- adian Field—Naturalist 90: 470-471. Maltby, L. S. 1978. Birds of the coastal zone of Melville Island, 1973-1975. Canadian Field-Naturalist 92: 24-29. 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pue purjsy syueg wo.J jst] p10de1 Surpse.1q/saisedg *(papnjauo2) xipusddy - Night Surveys of Yellow Rails, Coturnicops noveboracensis, and Virginia Rails, Rallus limicola, in Alberta using Call Playbacks DAVID R. C. PRESCOTT!, MICHAEL R. NORTON? and ISABELLE M. G. MICHAUD Alberta Conservation Association, 6th Floor, 9945 108th Street, Edmonton, Alberta TS5J 4M9 Canada \Present address: Alberta Sustainable Resource Development, Suite 404, 4911 — 51st Street, Red Deer, Alberta T4N 6V4 Canada 2Present address: Canadian Wildlife Service, #200, 4999 — 98th Avenue, Edmonton, Alberta T6B 2X3 Canada Prescott, David R. C., Michael R. Norton, and Isabelle M. G. Michaud. 2002. Night surveys of Yellow Rails, Coturnicops noveboracensis, and Virginia Rails, Rallus limicola, in Alberta using call playbacks. Canadian Field-Naturalist 116(3): 408-415. The Yellow Rail (Coturnicops noveboracensis) and Virginia Rail (Rallus limicola) are inconspicuous and apparently rare species in Alberta. Wide scale, targeted surveys have never been conducted for either species, and virtually nothing is known about their distribution, populations, or other traits in the province. We compiled historical records of Yellow and Virginia Rails and conducted nocturnal field surveys using call playback as a first step in clarifying the status of these species in the province. Sixty-two Yellow Rail and 66 Virginia Rail historical sites were collated, with the majority of records occurring in the central and south-central parts of the province. A total of 431 surveys were conducted at 404 sites between 17 May and 6 July 2000. These sites included 69.4% and 60.6% of historical sites for Yellow and Virginia rails, respectively. We encountered 42 unique Yellow Rails and 59 unique Virginia Rails during the surveys, with only 7.0% and 17.5% of visited historical sites being occupied. Fifteen new sites for Yellow Rails were found. Most notable was the pres- ence of 21 birds on Hay Lake in northwestern Alberta, which is a 450 km extension to the known breeding range. Twenty- three new sites for Virginia Rails were found. Playbacks were more effective at detecting Virginia Rails, as the number of spontaneously calling individuals was lower than was found for Yellow Rails (55.0% vs. 80.4%). Yellow Rails were most likely to be found in seasonal wetlands containing sedge, whereas Virginia Rails occupied a wide variety of semipermanent and permanent wetlands. Yellow Rails were most likely to be found when there was little or no moon, and during the mid- dle (darkest) part of the night. Virginia Rails were found significantly more often when the moon was more than half full and relatively unobscured by clouds. The efficiency of future surveys for these species could therefore be improved by focusing efforts during periods and conditions when birds are most likely to be detected. Key Words: Yellow Rail, Coturnicops noveboracensis, Virginia Rail, Rallus limicola, Alberta, call playback surveys, dis- tribution, status. Rails are inconspicuous and reclusive marsh birds that are more likely to be detected by their calls than to be seen (Godfrey 1986). Given that calling is most likely to occur during hours of darkness, it is not sur- prising that the life history and population status of many species is poorly understood (Eddleman et al. 1988). Evidence suggests that habitat loss and other pressures have led to the reduction of populations of a number of species in North America (Eddleman et al. 1988). Clarification of population status and a better understanding of the ecology of these species should therefore be a high priority to conservation biologists (Holliman 1977; Eddleman et al. 1988; Tacha et al. 1994). Three species of rail regularly occur in Alberta. The Sora (Porzana carolina) is found in most types of freshwater wetlands, and is a common breeding species throughout the province (Salt and Salt 1976; Semenchuk 1992). The Yellow Rail (Coturnicops noveboracensis) which inhabits sedge meadows with little or no standing water (Bookhout 1995) is far less common in Alberta. - The species is officially listed as being of “undeter- mined” status in the province due to a lack of infor- mation on population size and trend (Alberta Environment 2000*), and is further designated as being of “special concern” by the Committee on the Status of Endangered Species in Canada (COSEWIC) (Alvo and Robert 1999*, Anonymous 2000*). Alvo and Robert (1999*) documented 40 sites where birds have been known to occur in the province, and speculated that there may be at least 500 breeding pairs in Alberta. The Virginia Rail (Rallus limicola), which occupies a broad range of wetland types (Conway and Eddleman 1994; Conway 1995) is also considered to be of “undeter- mined” status in Alberta (Alberta Environment 2000*). However, the species is common in many parts of its range and is not listed by COSEWIC as being a species at risk in Canada (Anonymous 2000*). A comprehensive compilation of historical records has not previously been attempted in Alberta, but the species was found in only 24 loca- tions in the province during the Alberta Bird Atlas *See Documents Cited 408 2002 project in the late 1980s and early 1990s (Semenchuk 1992). Our current knowledge of the status of Yellow and Virginia rails in Alberta is poor (Pinel et al. 1991), and has been derived from a variety of observations collected over several decades. These data provide no information on the size and distribution of popu- lations occurring in the province during any single breeding season, nor on the persistence of popula- tions at any one site. Furthermore, there is much available habitat that has not been surveyed (Alvo and Robert 1999*), and it is likely that both species are more common than indicated by current informa- tion. In this study, we aimed to clarify the status of the Yellow and Virginia rails in Alberta by: (1) iden- tifying sites in the province where the species are known to have occurred; (2) conducting inventories at these sites to determine occupancy by individuals during the breeding season; and (3) locating new potential breeding sites through inventories in suit- able habitat. We also attempted to gain a better understanding of the habitat requirements and calling behavior of these species to increase effectiveness of future field studies in the province. Methods 1. Compilation of Historical Records We obtained many site records for Yellow Rails from Alvo and Robert (1999*). Records for Virginia Rails, and additional records for Yellow Rails were obtained from the Biodiversity/ Species Observation Database (maintained by Alberta Fish and Wildlife Division and the Alberta Conservation Association), the Alberta Bird Checklist Program and Alberta Bird Atlas database (both administered by the Federation of Alberta Naturalists), known published reports, museum collections, the Breeding Bird Survey database, and through requests for information pub- lished in several provincial publications and internet newsgroups. 2. Field Surveys Yellow Rails have been reported calling in Alberta between 26 May and 28 July, whereas Virginia Rails have been heard as early as 14 May in the extreme south of the province (Pinel et al. 1991). We therefore determined that surveys between 20 May and 25 July would have the best chance of detecting birds if present, and that surveys should begin in the south to capitalize on earlier arrival of birds in that area. Past studies have suggested that the probability of detecting either Yellow or Virginia Rails at an occupied site during a single visit is approximately 75% (Bart et al. 1984; Gibbs and Melvin 1993), which we considered to be reasonable for a reconnaissance-level survey. However, repeat visits were occasionally made to sites if logistically feasible. We attempted to visit most documented sites, although the specific locations of many sites PRESCOTT, NORTON, AND MICHAUD: NIGHT SURVEYS OF RAILS 409 were not well described in available documentation. This was especially true of older records, but also of records from the Alberta Bird Atlas database which were only compiled by 10 x 10 km? squares (Semen- chuk 1992). In these cases, we typically surveyed between two and five areas of suitable habitat within five kilometers of the expected location, or within the identified Atlas square. Sites scheduled for noc- turnal surveys were usually visited during daylight so that access points could be determined, and gener- al habitat variables recorded (see below). Daytime reconnaissance also allowed identification of new areas of potentially occupied habitat that could be visited during the night. Nocturnal surveys were conducted between sunset and sunrise, which is the period when rails are most likely to vocalize (Johnson and Dinsmore 1986; Manci and Rusch 1988; Gibbs and Melvin 1993: Bookhout 1995; Robert and Laporte 1997). To increase the probability of detection, playback of con- specific calls was used (Marion et al. 1981; Gibbs and Melvin 1993). Calls used in playbacks were assembled from commercially available sources (Peterson 1992; Brigham, undated), and delivered from portable tape players at a height of approxi- mately 1.5 m and a volume between 80 and 95 db. Although the calling and responsiveness of both Yellow and Virginia rails is apparently not greatly affected by weather (Bart et al. 1984; Gibbs and Melvin 1993), wind and precipitation undoubtedly affect detection distances (Bart et al. 1984). We therefore conducted surveys under a range of condi- tions, but avoided periods with winds > 20 km/hr and heavy rainfall. Surveys consisted of a three-minute listening period, followed by three, 20-second play- backs of Yellow Rail calls (“clicks”) separated by 20 seconds of silence, and three, 20-second broadcasts of Virginia Rail calls, also separated by 20 seconds of silence (two sets of descending “grunts” and one set of combined “kadic-kadic” and “kicker” calls, which are the vocalizations most often used in territorial advertisement and mate attraction; see Kaufmann 1983; Conway 1995). A final three-minute listening period followed the playback sequence. In a few cases (< 10%), playback of only one species’ call was delivered (generally at sites historically occupied by that species, and with no obvious habitat for the other species). The number of birds calling during each of the pre-playback, playback and post-playback was recorded. We recorded air temperature (°C), wind speed (Beaufort scale), the presence of precipitation or fog, cloud cover (%), moon phase (new, < half full, > half full, or full), moon visibility (visible, obscured, or absent) and time of night (early = 22:00 to 23:59 h; middle = 24:00 to 01:59 h; late = 02:00 to 05:30 h) as factors that may influence calling behav- ior. We also recorded (usually during daylight hours, see above) the dominant vegetation type at the survey 410 location (cattail, bulrush, sedge, or mixed), and the approximate percentage of major cover types in the wetland (standing emergents, open water, bare ground, shrubs, or trees). In addition, we recorded whether surveyed wetlands were permanent (water present in all years) or semipermanent (dry in some years) based on descriptions on topographic maps, or seasonal (present for only part of the year). Attributes of locations where each species was detected were compared with sites where individuals were appar- ently absent using Mann-Whitney U-tests or chi- square analyses (Conover 1980). Results I. Compilation of Historical Records We compiled a total of 62 historical sites for Yellow Rails and 66 sites for Virginia Rails in the province, which encompassed at least 95 individual records for Yellow Rails reported between 1914 and 2000, and 90 records for Virginia Rails observed between 1945 and 1999. Although the specific dates of some records were unclear, one location for Yellow Rails and at least five locations for Virginia Rails were known only from individuals encountered on or before 1 May, or later than 31 August. Al- FiGuRE 1. Distribution of historical records for Yellow (A) and Virginia (B) Rails in Alberta. See Prescott et al. (2001*) for details. Solid symbols indicate sites that were surveyed using call playbacks during 2000. THE CANADIAN FIELD-NATURALIST Vol. 116 - though there is no information on the timing of migration by these species in Alberta (Pinel et al. 1991), we consider these records to be questionable in terms of identifying historical breeding sites. Overall, the data show that both species have been most commonly encountered in the central and south-central parts of the province, with the range of the Virginia Rail extending slightly further to the southeast, and the Yellow Rail to the northeast (Figure 1). Complete details on individual site records are available in Prescott et al. (2001*). 2. Field Surveys A total of 431 surveys were conducted between 17 May and 6 July 2000. This total included 17 sites that were visited twice, and five sites that were visit- ed on three occasions during the survey period. A total of 404 unique sites were surveyed, covering most of the province with the exception of the mountain parks and extreme northern and northeast- ern areas (Figure 2). Calls of Yellow and Virginia rails were played during 398 (92.3%) and 414 (96.1%) surveys, respectively. Forty-three of the 62 (69.4%) historical sites for Yellow Rails, and 40 of the 66 (60.6%) known Virginia Rail sites were visit- ed and checked for the presence of potential habitat. 2002 A Both Species A Virginia Only © Yellow Only FIGURE 2. Distribution of 404 sites surveyed for calling Yellow and Virginia Rails during 2000. Symbols indicate which call playbacks were delivered at each location. No suitable habitat could be found at six historical sites (three for each species), so no playback surveys were completed at these locations. Using only the maximum number of birds heard over all visits to a particular site, we estimate that 42 unique Yellow Rails and 59 unique Virginia Rails were encountered at 18 and 35 sites, respectively. Only three (7.0%) of the visited historical Yellow Rail sites, and seven (17.5%) of the Virginia Rail sites were found to support a total of 5 and 19 indi- viduals, respectively (Tables 1 and 2). However, a PRESCOTT, NORTON, AND MICHAUD: NIGHT SURVEYS OF RAILS Ai] minimum of 37 additional Yellow Rails were detect- ed at 15 sites where the species was not previously known to occur. This included 21 birds found at six sites on Hay Lake in extreme northwestern Alberta, about 450 km NNW of the nearest historical location for the species (Kimiwan Lake). We also found an estimated 39 Virginia Rails at 23 sites (24 surveys) at non-historical locations. Only one site was found to contain both species. The geographical distribution of rails found during the 2000 survey shows a concentration of sites for Virginia Rails in central and southern Alberta (with the exception of one record near La Crete in north- western Alberta), whereas the Yellow Rail was sparsely and widely distributed from central to northwestern Alberta (Figure 3). 3. Influences on Calling Behavior and Distribution Of 46 Yellow Rails detected on field surveys (including multiple visits to sites), 37 (80.4%) were calling spontaneously (16 of 20 sites), with an addi- tional nine individuals responding to tape playback. Six of these individuals responded quickly (i.e., dur- ing playback), with an additional three birds respon- ding only during the post-playback period. The pres- ence of Yellow Rails at four sites was determined only because of the use of playbacks. At three sites however, birds calling spontaneously stopped calling immediately when tapes were played, and did not start calling again before the session was terminated. Detection of Yellow Rails was influenced by the lunar phase, and birds were most vocal when illumination levels were low. Specifically, birds were found at 12.2% of 49 sites surveyed during a new moon, 4.4% (n = 159) and 2.6 % (n= 114) of sites when the moon was less than and more than half full, respectively, and only 1.4 % (n=71) of sites visited during a full TABLE 1. Location and notes on where Yellow Rails were detected during the 2000 survey. Survey Location! Latitude ((N) Longitude (°W) Notes 22 km W of Airdrie 51325 114.328 2 birds on 2 July Lochend Lake 51.330 114.321 1 bird on 2 July 5.5 km NE Rocky Mountain House 52.327 114.954 2 birds on 3 July 2.5 km W of Mirror 52.463 113.164 1 bird on 17 June Spotted Lake (*) 52.487 113.133 3 birds on 31 May, | bird on 4 June (daytime) Reflex Lake (*) 52.660 110.027 1 bird on 22 June 17 km WSW of Edson on Hwy 16 53.543 116.668 2 birds on 28 June 1.5 km NE of Angling Lake 54.228 110.204 4 birds on 22 June, 3 birds on 23 June 7.2 km NNE of Slave Lake (*) 55.346 114.726 1 bird on 30 June. Also 2 Virginia Rails at this site Buffalo Bay 55.563 116.273 1 bird on 30 June 198 km N of Red Earth 58.049 115.612 2 birds on 6 July Lost Lake 58.345 116.402 | bird on 5 July NE Shore of Hay Lake 58.833 118.633 | bird on 5 July 58.832 118.628 3 birds on 5 July 58.832 118.621 5 birds on 5 July 58.833 118.648 5 birds on 5 July 58.826 118.697 | bird on 5 July 58.864 118.871 6 birds on 5 July *historical site listed in Prescott et al. (2001*) Re err RIES 412 THE CANADIAN FIELD-NATURALIST Vol. 116 aa TABLE 2. Location and notes on where Virginia Rails were detected during the 2000 survey. Survey Location! Latitude (°N) Longitude (°W) Notes Prince’s Spring (*) 50.813 110.344 1 bird on 30 May 50.808 110.341 1 bird on 30 May 6.5 km S of Priddis 50.848 114.325 1 bird on 5 June 15 km NNE of Oyen 51.472 110.355 2 birds on 14 June 0.5 km N of Barrie Lake (*) 51.766 114.191 3 birds on 16 June 51.780 114.193 4 birds on 16 June North of Barrie Lake 31783 114.191 1 bird on 16 June 15 km S of Delburne (*) 52.061 113.279 1 bird on 16 June 11 km WSW of Lousana 52.095 113.338 1 bird on 16 June 3.5 km SE of Alix 52)388 113.145 4 birds on 16 June 6 km E of Alix 52.405 113.096 1 bird on 17 June 0.8 km E of Gabriel Lake 52.426 114.410 1 bird on 19 June 2 km NW of Lacombe (*) 52.485 113.764 1 bird on 17 June 52.491 113.749 1 bird on 17 June Ribstone Creek 52.488 110.740 3 birds on 21 June 6 km W of Chauvin 52.700 110.221 2 birds on 21 June 1 km W of Ribstone 52722 110.281 1 bird on 21 June 1 km SW of Samson Lake (*) S22 £13.262 3 birds on 19 June a2 lA: 113.282 1 bird on 19 June 52.720 113.284 2 birds on 19 June Bearhills Lake 52.960 113.626 1 bird on 19 June. Not heard on previous night 2 km W of Camrose 53.018 112.881 2 birds on 22 June 8 km NW of Camrose 53.0055 112.953 1 bird on 22 June 1 km NE of Bittern Lake village 53-105 113.028 2 birds on 23 June 8 km N of Camrose 53.109 112.825 1 bird on 27 June 21 km NE of Camrose 53-1935 112.680 1 bird on 27 June 5.5 km NW of Big Hay Lake 53.234 113.221 1 bird on 22 June 8 km W of Mannville Pe hora 111.305 2 birds on 22 June Tawayik Lake Marsh, Elk Island National Park 53.618 112.874 2 birds on 26 June Elk Island National Park, S of N gate (*) 53.700 112.809 1 bird on 22 June 1.8 km NE of Angling Lake 54.241 110.204 1 bird on 22 June, 3 birds on 23 June 0.5 km S of Cherry Grove 54.332 110.079 3 birds on 21 June 7.2 km NNE of Slave Lake 55.346 114.726 2 birds on 30 June. Also 1 Yellow Rail at this site S307, 114.752 1 bird on 30 June 3.5 km SW of La Crete (*) 58.155 116.376 1 bird on 5 July *historical site listed in Prescott et al. (2001*) moon (x?2= 9.7, df = 3, p < 0.05). Rails were also found more frequently during the middle of the night (24:00-01:59 h) than during earlier or later periods (x?= 8.2, df = 2, p< 0.05). Although these patterns suggest that darkness is a factor in calling behavior, detection was not influenced by whether the moon was visible, obscured or absent during a survey (x?= 0.31, df =2, p>0.8). There was also no appar- ent influence of any other meteorological variables (wind, precipitation, cloud, fog or temperature; all p > 0.4) on encounters with Yellow Rails. Spontaneously-calling Virginia Rails were detect- ed at 23 sites (total of 33 birds, or 55.0% of all encounters). Responses to tapes were generally stronger than for Yellow Rails. An additional 27 birds were detected through playbacks, with 22 of these responding immediately. Thirteen sites for Virginia Rails were identified only through respons- es to playbacks. At five sites, however, birds (six individuals) that were already vocalizing became quiet upon delivery of the playback. Calling by Virginia Rails was influenced by the moon phase (x2 = 11.2, df = 3, p < 0.01), but in the opposite pat- tern to Yellow Rails. Birds were most likely to be found during a full moon (13.9 % of 72 sites sur- veyed) or periods when the moon was more than half full (13.5% of 159 sites), and least likely when the moon was less than half full (4.4% of 126 sites) or during a new moon (3.9% of 52 sites). Birds also were detected more often on nights when the moon was visible or obscured, as opposed to being absent (x?= 10.2, df = 2, p< 0.01). Cloud cover, precipita- tion and time of night did not differ between sites where Virginia Rails were considered present or absent, although temperatures during surveys when birds were detected were lower than when no birds were found (8.8 + 0.6°C vs. 10,0:+.0.2°C; Taz p < 0.05). 2002 PRESCOTT, NORTON, AND MICHAUD: NIGHT SURVEYS OF RAILS 413 O Yellow Rail A Virginia Rail FIGURE 3. Sites where Yellow and Virginia Rails were found during call playback surveys in 2000. See Tables 1 and 2 for additional details. Yellow Rails were detected significantly more often than expected in sedge and “mixed” (cattail/ bulrush/sedge) wetlands, and less often in basins dominated by cattail or bulrush (x?= 8.4, df =3, p < 0.05). Furthermore, occupied basins were most likely to be seasonal, as opposed to semipermanent or permanent (x?= 6.7, df = 2, p< 0.05), and tended to have less open water (10.6 + 4.4 [SE]%) than sites where birds were not detected (24.8 + 1.5%; T = 2.8, p < 0.005). Virginia Rails showed no preference for basins dominated by particular vegetation types (p > 0.4), but preferred wetlands with more open eee 45% ys. 23.5. 246%; T=2.2, p < 0.05) and less shrub (5.1 + 1.3 vs. 9.9 + 0.8; T = 1.9, p< 0.06). Furthermore, birds were found more often on permanent or semipermanent wetlands than on seasonal waterbodies (x2= 5.9, df = 2, p < 0.06). Discussion Our results support previous descriptions of Yellow and Virginia rails being widely distributed but relatively uncommon species in Alberta (Salt and Salt 1976; Pinel et al. 1991; Semenchuk 1992; McGillivray and Semenchuk 1998). In general, these references identify the central areas of the province as being the areas of highest density, which is con- sistent with the distribution of historical records that we compiled. We caution, however, that higher num- bers of human observers and higher road density likely bias the distribution of historical records in that part of the province, and that the species’ histor- ical occurrence in other parts of the province may be underestimated. We focused our field surveys around historical sites, and therefore many of the previously unsur- veyed areas tended to be near, or between historical locations. It is therefore not surprising that the majority of encounters with rails (and especially Virginia Rails) in 2000 occurred in the central part of the province as well. The distribution of Yellow Rails was less closely matched to historical records, with field surveys revealing a relatively sparse distri- bution from central to northwestern Alberta. The records in the northwest were especially notable, as they represent a major extension of the known range of this species. In fact, almost half of all Yellow Rail encounters in 2000 were at six sites on Hay Lake. Those sites represent a very small fraction of the available habitat in the Hay-Zama Lakes area (K. Wright, Alberta Conservation Association, personal communication), and the region has the potential to Support many more birds than we encountered. This is also true for the northeastern part of the province, which has poor road access and was not well sur- veyed in 2000. Although Virginia Rails have never been reported from northeastern Alberta, there are several historical records of Yellow Rails from the area north of Fort McMurray, and from adjacent areas of the Northwest Territories (see Alvo and Robert 1999*). The Peace-Athabasca Delta, which has one historical Yellow Rail record, contains large areas of wet sedge (M. Bradley, Parks Canada, per- sonal communication) and has the potential to sup- port large numbers of breeding rails. We also did not survey potential habitat in Jasper and Banff National Parks. These areas have historical records for both Yellow and Virginia rails, although none of these occurred with certainty during the breeding season (Prescott et al. 2001*). However, there are several records of both species during the breeding season from adjacent areas of British Columbia (Campbell et al. 1990), which suggests that the mountainous areas of Alberta should be considered for surveys in the future. Of approximately 400 surveys conducted for each species, we found Yellow and Virginia rails at only 18 and 35 sites, respectively. Furthermore, 12 (66.7%) of the Yellow Rail sites and 29 (82.9%) of the Virginia Rail sites contained only a single calling bird (Tables 1 and 2). Values for both the number of sites occupied and the number of calling birds at each site are likely underestimates, as > 90% of all the sites were only visited on a single occasion which would have detected occupancy with no better than 75% certainty (Bart et al. 1984; Gibbs and Melvin 1993). Nevertheless, it is apparent that both species of rails are relatively uncommon within their Alberta range. Occupancy rates at historical sites were also low. This is especially true of Yellow Rails (three of 42 sites surveyed), and may reflect the observed preference of birds for seasonal wet- lands ({[see also Boukhout (1995) and references therein)] which can vary greatly in condition from 414 year to year depending on local precipitation. Return rates by breeding birds might therefore be even lower than the 5.4% reported by Robert and Laporte (1999) for relatively stable estuarine habitats in east- ern Quebec. Seasonal wetlands on the prairies are also most likely to be impacted by agricultural activ- ities such as cultivation, drainage, and cattle grazing (several historical areas visited in 2000 were dry, and being grazed). In contrast, Virginia Rails were found most often in permanent or semipermanent wetlands, where habitat suitability is more stable over time, and which are less likely to be altered by human activities. Evidence suggests that Virginia Rails will return to sites in successive years if habitat conditions remain stable (Mousley 1931; Conway 1995), and the higher occupancy of historical sites we observed (seven of 39) may reflect this stability. The relatively low occupancy rate of both species suggests caution must be exercised in using compila- tions of historical data collected over many years to assess population status of these species (e.g., Alvo and Robert 1999*). Attempts to inventory Yellow and Virginia rails in Alberta have been scant (Pinel et al. 1991; Semenchuk 1992), and our surveys represent the first focused effort to determine the distribution and sta- tus of these species in the province. However, there remain several key areas of the province that were not surveyed, and much suitable habitat in all parts of the province that could not be visited. Clarification of the status of these species therefore demands additional survey effort from biologists. However, we recommend that additional large-scale inventories be preceded by research into factors affecting calling behavior of these two species. Of particular interest is our observation that Yellow Rails tended to be found during periods with little or no moon (and during the darkest part of the night), and that Virginia Rails tended to be found during periods where the moon was more than half full and the sky was relatively clear. Although we hesitate to place strong emphasis on patterns resulting from an observational study with a relatively small sample of occupied sites, the results suggest that the survey protocol could be refined to focus efforts during periods when birds are most likely to be detected. Furthermore, if the probability of detecting sponta- neously-calling birds could be increased, the use of call playbacks could be eliminated. This would reduce the amount of time required to inventory each area, and allow many more locations to be surveyed in a single field season. Our surveys in 2000 have identified a number of sites where season-long moni- toring of calling behavior could be conducted, and we intend to initiate intensive surveillance of select- ed sites in 2001. THE CANADIAN FIELD-NATURALIST Vol. 116 - Acknowledgments We acknowledge the dedicated field assistance of M. Piorecky, C. Morcos, J. Adamyk, L. Engley, R. Schmelzeisen, G. McLelland, S. Feser, R. Corrigan, D. Fairless, K. Wright, K. Morton, M. Vonhof and D. Eckford, and the numerous biologists and naturalists who offered information on historical site locations or areas of potential habitat. We also thank W. Hawkins (Ducks Unlimited Canada) for providing remote sens- ing data, B. Dale (Canadian Wildlife Service) for gen- erous logistical support, T. Wiens (Federation of Alberta Naturalists) for supplying Alberta Bird Atlas data, and A. J. Erskine and M. Robert for comments on an earlier draft of the manuscript. The study received financial assistance from the North American Waterfowl Management Plan, Canadian Wildlife Service, Alberta Conservation Association and Alberta Sport, Recreation, Parks and Wildlife Foundation. Alberta Environment (Natural Resources Service) provided logistical support. Documents Cited [marked * in text citations] Alberta Environment. 2000. Status of Alberta wildlife 2000. Preliminary status evaluation of the birds: non- passerines. Alberta Environment, Fisheries and Wildlife Management Division, Edmonton, Alberta. 182 pages. Alvo, R., and M. Robert. 1999. COSEWIC status report on Yellow Rail (Coturnicops noveboracensis). Commit- tee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 56 pages. Anonymous. 2000. Canadian species at risk. Committee on the Status of Endangered Wildlife in Canada, Ottawa, Ontario. 19 pages. Prescott, D. R. C., M. R. Norton, and I. M. G. Michaud. 2001. A survey of Yellow and Virginia Rails in Alberta using nocturnal call playbacks. Alberta Conservation Association, Edmonton, Alberta. 20 pages. Literature Cited Bart, J., R. A. Stehn, J. A. Herrick, N. A. Heaslip, T. A. Bookhout, and J. R. Stenzel. 1984. Survey methods for breeding Yellow Rails. Journal of Wildlife Manage- ment 48: 1382-1386. Boukhout, T. A. 1995. Yellow Rail (Coturnicops novebo- racensis). Number 139 in The birds of North America. Edited by A. Poole and F. Gill. The Academy of Natural Sciences, Philadelphia, PA, and The American Ornithol- ogists’ Union, Washington, D.C. 16 pages. Brigham, M. [Undated]. Bird sounds of Canada. Volume 1: Loons to woodpeckers. Great Wildlife Recordings, Manotick, Ontario. Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. W. Kaiser, and M. C. E. MecNall. 1990. The birds of British Columbia. Volume 2: Nonpasserines. Royal British Columbia Museum, Vic- toria, British Columbia. Conover, W. J. 1980. Practical nonparametric statistics. John Wiley and Sons, New York, NY. 493 pages. Conway, C. J. 1995. Virginia Rail. Number 173 in The birds of North America. Edited by A. Poole and F. Gill. 2002 The Academy of Natural Sciences, Philadelphia, Pen- nsylvania, and The American Ornithologists’ Union, Washington, D.C. 20 pages. Conway, C. J., and W. R. Eddleman. 1994. Virginia Rail. Pages 193-206 in Management of migratory shore and upland game birds in North America. Edited by T. C. Tacha and C. E. Braun. International Association of Fish and Wildlife Agencies, Washington, D.C. Eddleman, W. R., F. L. Knopf, B. Meanley, F. A. Reid, and R. Zembal. 1988. Conservation of North Ameri- can rallids. Wilson Bulletin 100: 458-475. Gibbs, J. P., and S. M. Melvin. 1993. Call-response sur- veys for monitoring breeding waterbirds. Journal of Wildlife Management 57: 27-34. Godfrey, W. E. 1986. The birds of Canada. Revised edi- tion. National Museums of Canada, Ottawa, Ontario. 595 pages. Holliman, D. C. 1977. Rails and gallinules. Pages 118- 121 in Management of migratory shore and upland game birds in North America. Edited by G. C. Sanderson. Internattional Association of Fish and Wildlife Agencies, Washington, D.C. Johnson, R. R., and J. J. Dinsmore. 1986. The use of tape-recorded calls to count Virginia Rails and Soras. Wilson Bulletin 98: 303-306. Kaufmann, G. W. 1983. Displays and vocalizations of the Sora and the Virginia Rail. Wilson Bulletin 95: 42-59. Manci, K. M., and D. H. Rusch. 1988. Indices to distri- bution and abundance of some inconspicuous waterbirds on Horicon Marsh. Journal of Field Ornithology 59: 67-75. PRESCOTT, NORTON, AND MICHAUD: NIGHT SURVEYS OF RAILS 415 Marion, W. R., T. E. O’Meara, and D. S. Maehr. 1981. Use of playback recordings in sampling elusive or secre- tive birds. Studies in Avian Biology 6: 81-85. McGillivray, W. B., and G. P. Semenchuk. 1998. Field guide to Alberta birds. Federation of Alberta Naturalists, Edmonton, Alberta. 350 pages. Mousley, H. 1931. Notes on the home life of the Virginia Rail. Canadian Field-Naturalist 45: 65-66. Peterson, R. T. 1992. A field guide to western bird songs, 2nd edition Houghton-Mifflin, New York, NY. Pinel, H. W., W. W. Smith, and C. R. Wershler. 1991. Alberta Birds, 1971-1980. Volume 1: Non-passerines. Provincial Museum of Alberta Natural History Occa- sional Paper Number 13. 243 pages. Robert, M., and P. Laporte. 1997. Field techniques for studying breeding Yellow Rails. Journal of Field Orni- thology 68: 56-63. Robert, M., and P. Laporte. 1999. Numbers and move- ments of Yellow Rails along the St. Lawrence River, Quebec. Condor 101: 667-671. Salt, W.R., and J. R. Salt. 1976. The birds of Alberta. Hurtig Publishers, Edmonton, Alberta. 498 pages. Semenchuk, G. P. Editor. 1992. The atlas of breeding birds of Alberta. Federation of Alberta Naturalists, Edmonton, Alberta. 391 pages. Tacha, T. C., C. E. Braun, and R. E. Tomlinson. 1994. Migratory shore and upland game bird resources — sta- tus and needs. Pages 219-223 in Migratory shore and upland game bird management in North America. Edited by T. C. Sacha and C. E. Braun. International Association of Fish and Wildlife Agencies, Washington, D.C. Received 11 May 2001 Accepted 28 June 2002 The Gray Wolves, Canis lupus, of British Columbia’s Central and North Coast: Distribution and Conservation Assessment Curis T. DARIMONT! and PAUL C. PAQUET?: 3 \Department of Biology, University of Victoria. PO Box 3020, Victoria, British Columbia V8W 3N5 Canada 2World Wildlife Fund Canada. 245 Eglinton Avenue East, Suite 410, Toronto, Ontario M4P 3J1 Canada 3Faculty of Environmental Design, University of Calgary, 2500 University Drive Northwest, Calgary, Alberta T2N 1N4, Canada Darimont, Chris T., and Paul C. Paquet. 2002. The Gray Wolves, Canis lupus, of British Columbia’s central and north coast: Distribution and conservation assessment. Canadian Field Naturalist 116(3): 416-422. The Gray Wolves (Canis lupus) of coastal British Columbia are a remnant group of a much larger population that once inhabited most of North America, including its west coast temperate rainforests. During summers 2000 and 2001, we sur- veyed 36 islands and 42 mainland watersheds on British Columbia’s Central and North Coast for the presence of wolves. An extensive survey had not been conducted previously. We observed wolf sign at all locations, including islands or island groups separated by approximately 7, 8, and 12-km from other large landmasses. The distribution of wolves on islands may be dynamic, with occupancy by solitary Wolves or packs being ephemeral. The potential for an island to support a persis- tent population of wolves may depend on the presence and abundance of their main prey, Black-tailed Deer (Odocoileus hemionus), and security from exploitation by humans. These factors likely are mediated by island isolation, area, shape, topography, and extent of logging. Mounting evidence suggests that logging negatively affects Wolves in temperate rain- forests by reducing carrying capacity for deer. Key Words: Gray Wolf, Canis lupus, distribution, islands, British Columbia, logging, conservation. The Gray Wolf, Canis lupus, is one of the most intensely studied wildlife species in North America, yet complete and recent distributional data are absent from the large and remote Central and North Coast of British Columbia (Figure 1). The equilibri- um theory of island biogeography predicts that species composition on islands is a function of immigration and extinction, mediated by island area and isolation (MacArthur and Wilson 1967). The ability of a terrestrial mammal to colonize islands depends on degree of isolation, the species’ swim- ming ability, and water conditions. Earlier, it was postulated that the occurrence of Gray Wolves on the islands of coastal British Columbia and Alaska was a function of the presence of their main prey, Black-tailed Deer (Odocoileus hemionus), but limited by isolation on islands to which wolves were capable of swimming (McCabe and Cowan 1945; Klein 1965). MacDonald and Cook (1996) stated that Gray Wolves occurred in all mainland areas and on most islands of southeast Alaska, but not on small and isolated ones without adequate prey. Nagorsen (1990) described Gray Wolf range in British Columbia as comprising the “entire mainland, Vancouver Island and some adja- cent islands”. Our main objective herein is to describe the distribution of Gray Wolves along the Central and North Coast of British Columbia, partic- ularly on islands. We also discuss conservation con- cerns relevant to wolves of coastal temperate rain- ° forests because these factors may influence future distribution (Person et al. 1996*; Darimont and Paquet 2000*; Person 2000). We recorded the infor- mation while conducting field studies of wolves and their prey during summers 2000 and 2001 (Darimont and Paquet 2000*; Darimont and Reimchen 2003; Darimont et al. in press). Methods Study Area Boats and airplanes provide the only access to this nearly roadless and mostly unsettled region. Extensive fjords divide mainland valleys. Tidal waters separate islands that vary from < one km? to > 2,200 km2 (Princess Royal Island). Inter-island and mainland-island distances range from several metres to approximately 13-km. The study area was roughly delineated by the Kshwan Valley (55° 37’ N, 129° 48’ W) in the north to Cape Caution (51° 10’ N, 127° 47' W) in the south, and oriented parallel to the coastline (Figure 1). The Coast Mountains and the Pacific Ocean bound the study area to the east and west respectively. Most of the low elevation forest is within the Coastal Western Hemlock biogeoclimatic zone (sensu Krajina 1965), characterised by a wet and temperate climate. Annual precipitation exceeds 350 cm in most areas. Thirty-year average annual snowfall measured near sea level varies from 86 cm (Bella Bella) to 155 cm (Ocean Falls) (Environment Canada 1991*). *See Documents Cited section. 416 | : | : 2002 DARIMONT AND PAQUET: WOLVES OF BRITISH COLUMBIA’S COAST 417 Pacific Ocean 19 McCauley Isiand 20 enecon Iniet River y ' -M) : 42 Campania Island : | . 5 43 Klekane River nS AON 44 Aaltanhash River } 45 Khutze River : I Lak 55 Naples River 56 Ellersi 78 Takush River FiGuRE |. Study area, survey sites, and other notable landmarks relevant to survey for Gray Wolf distribution on British Columbia’s Central and North Coasts during summers 2000 and 2001. Snowfall is much greater in the inland and higher ele- tailed Deer, Beaver (Castor canadensis), River Otter vation portions of the study area but no weather data = (Lutra canadensis), other mustelids, birds, and small exist. Prey species available to wolves include Black- _ rodents. Moose (Alces alces) inhabit the eastern fringes 418 of the study area. Mountain Goats (Oreamnos ameri- canus) are found on the mainland, but presence has been confirmed on only Pitt and Princess Royal Islands (Nagorsen and Keddie 2000). Marine foods such as spawning salmonids (Darimont and Reimchen 2002; Darimont et al. in press, and Paquet, unpublished) and beached marine mammals are also available. Possible competitors are Brown Bear (Ursus arctos) and Black Bear (U. americanus), Wolverine (Gulo gulo), Coyote (Canis latrans), and Cougar (Felis concolor). Environmental conditions that may influence movements of wolves among landmasses vary sea- sonally and geographically. The following informa- tion is summarised from Thomson (1981). Mean air and water temperatures are lower during winter than summer, and average wind speed and wave height are greatest during fall and winter and in outer coastal areas. Surface currents flood primarily to the northeast and ebb to the southwest, but are consis- tently modified by wind, runoff, bathymetry and shoreline configuration. Typically, current velocity increases in narrow waterways. Fog is most common during summer and on the outer coast. We used a Geographic Information System (ArcView 3.2 — Environmental Systems Research Institute Inc.) to calculate island areas from 1:20 000 forest inventory maps (Province of British Colum- bia) generalized to 1:250 000. Minimum shore-to- shore distances at low tides were calculated from marine charts (Canadian Hydrographic Service). We indexed isolation as the shortest island-to-mainland distance or sum of island-to-island distances (exclud- ing distances across islands) to mainland for outer islands (Conroy et al. 1999). We also measured min- imum distances to other large landmasses (>75- km). We used 75 km? as a threshold area because that is about the size of Coronation Island, southeast Alaska, on which some wolves among an initial introduced population of four survived in isolation for eight years (Klein 1996). Sampling We selected sampling sites that were well dis- tributed throughout the study area and where we considered moorage safe. At each location, we used sandy beaches, estuaries, and forests of the beach fringe to begin our search for wolf sign. Wildlife trails, often next to watercourses, allowed travel inland. We also surveyed logging roads when encountered and often circumnavigated Beaver ponds and other wetlands. In addition, we walked forest ridgelines. Our surveys rarely extended greater than 5 km inland. Survey effort differed at each loca- tion from a few hours to several days. We determined presence of wolves by noting tracks, scat, sightings, and carcasses. We did not sys- tematically solicit howling but recorded all vocaliza- tions. Wolves are the only canid known on coastal islands in the region, although Red Foxes (Vulpes vulpes) and Coyotes occur on the mainland THE CANADIAN FIELD-NATURALIST Vol. 116 (Nagorsen 1990). Coyote tracks can be distinguished from wolf tracks based on differences in size (Rezendes 1999). Scats from Coyotes are generally smaller than scats from wolves, although there is some overlap in size (Weaver and Fritts 1979). Although the probability in this remote area is low, we may have assigned Gray Wolf presence to the tracks or scats of large feral Dogs (Canis familiaris). Results We observed recent wolf sign in all mainland watersheds (n = 42), confirming a continuous north- south distribution on the coastal mainland. Thirty- four of 36 islands had recent Gray Wolf sign. The most isolated islands on which we noted Gray Wolves were Moore Island, the Goose Group, and adjacent Dundas/Dunira Islands at 5 km, 7 km, and 12/13 km respectively from other large landmasses (Figure 1; Table 1). We found scats and tracks at all survey locations (n = 78) except the Goose Group and Moore Island, where no tracks were observed (Table 1). We found remains of dead wolves at three island sites and observed wolves on the shoreline at several main- land rivers and on six islands (Table 1). Howling was heard at some sites (Table 1). On the Goose Group of islands (~25 km2), we observed extensive sign of deer and found one old scat from a wolf. A Steller Sea Lion (Eumetopias jubatus) carcass showed no evidence of scavenging by large mammals. On the Moore Islands, which comprise a very small (<5 km?) archipelago, we recorded no sign of deer. However, we found one old scat containing bird and other unidentified mate- rial. In contrast, we noted deer sign and fresh wolf tracks and scat on Dundas and Dunira Islands, which are relatively large but extremely isolated adjacent islands at least 12 km from other large (or any habit- able) landmasses (Table 1). Discussion The current distribution of wolves in coastal British Columbia ranges from the deepest inlets of the coastal mountains to the most remote of islands (Figure 1; Table 1). The presence of wolves at all mainland survey sites concurs with earlier reports of the species’ distribution along the Pacific Coast (Nagorsen 1990; MacDonald and Cook 1996). Notably, of the 17 islands surveyed by both stud- ies, we detected the presence of wolves on six islands that McCabe and Cowan (1945) did not. These dis- crepancies likely reflect an artifact of sampling effort (ours being greater and more focused), or perhaps (re)colonization. Wolf and deer sign was detected on the Goose Group, where Guiguet (1953), after com- pleting a four-month ecological inventory, reported neither species. Deer may have colonized the group since then (either naturally or assisted by humans). 2002 DARIMONT AND PAQUET: WOLVES OF BRITISH COLUMBIA’S COAST 419 TABLE I. Island areas (km?) and distances (m) from mainland and islands greater than 75 km? for island sampling sites on the Central and North Coast of British Columbia, Canada where Gray Wolves or their sign were observed. ca = carcass, ho = howling, ob(x,y) = observation (number seen at x location and at a different y location), sc = scat, tr = tracks. N/A = not applicable (i.e., island is closer to mainland than other landmasses). Sampling site codes match those in Figure 1. Sampling Sign Island Site Observed Pearse 9 ob(3),sc,tr Wales 11 sc,tr Dundas 13 sc, tr Dunira 15 sc, tr Stephens 16 sc,tr Lewis 17 ob(2),sc,tr Porcher 18 ‘sc,tr McCauley 19 sc,tr Anger 26 ho,ob(1),sc,tr Hawkesbury 29 ca,ho,ob(2),sc,tr Banks 30 sc,tr Pitt 31 ho,ob(3,1),sc,tr Farrant 32 ho,ob(4,1),sc,tr Gribbell 33 sc,tr Fin 38 sc,tr Gil 40 sc,tr Dewdney 4] sc, tr Compania 42 sc,tr Princess Royal 47 ca,sc,tr Moore 49 sc Aristazabal 50 sc,tr Pooley 51 sc, tr Roderick 53 sc,tr Yeo 59 ob(1),sc,tr Chatfield 60 sc,tr Athlone 62 sc,tr Dufferin 63 sc,tr Horsfall 64 sc,tr Cunningham 65 sc,tr Stryker 66 sc, tr Campbell 67 ca,sc,tr Denny 68 sc,tr King 69 sc,tr Goose Group 70 sc Hunter Vg ob(1),sc,tr Calvert 73 sc,tr The presence of wolves on extremely isolated islands suggests that the potential for wolves to swim among landmasses is high. The ocean, however, likely serves as a considerable barrier to movement. Pre- viously, Gray Wolves have been described as good swimmers, but only of short distances in fresh water (Mech 1970; Pimlott et al. 1969*; Nelson and Mech 1984; Coscia 1993). Swimming in marine environ- ments probably carries considerably more energetic costs than travel on land, and may impose costs by elimination (e.g., drowning). Moreover, the effects of currents, which vary between landmasses, may be as important as straight-line distances in influencing movement (Cameron 1958; Williamson 1981). Finally, sightability [visibility] may affect orientation (MacArthur and Wilson 1967) and influence which Distance Distance Area to to Island (km2) Mainland (m) > 75 km? (m) 226 300 N/A 97 700 350 160 12000 N/A 22 13050 1700 78 3100 700 7 2400 100 632 2300 1900 273 1050 650 mW | 600 200 322 1950 1550 1024 3250 2200 1349 400 N/A 50 450 50 207 1550 1550 13 2250 1750 238 3900 2050 37 6200 100 157 2450 2050 2295 900 50 5 12500 8000 451 4350 3450 162 250 100 239 350 100 95 250 N/A 48 1000 100 40 2800 1350 43 2750 1300 32 2650 1250 115 850 150 10 2800 1400 145 1450 400 177 1000 150 826 1750 N/A 24 8700 7250 399 1700 700 326 3850 3100 islands wolves select, especially during foggy periods of summer. Radio-telemetry data from southeast Alaska con- firm that dispersal across large water bodies is possi- ble but infrequent. Person (2000) documented wolves swimming at least 2 km, although none of 11 dispersing animals left Prince of Wales or adjacent islands during three years of study. Although low prey density can stimulate dispersal (Peterson and Page 1988; Fuller 1989), presumably starving wolves on Coronation Island, southeast Alaska, failed to swim 900 m to a nearby island where deer were available (Klein 1996). Due to the barrier effects of water, some popula- tions on islands are probably independent sub-popu- lations among which exchanges of individuals or 420 packs are limited. We were not able to determine if the wolf sign found on islands came from multiple or solitary animals, or whether Wolves were resident or transient. Future studies may benefit by examining the presence, movements, and demographic fates of individuals over time to examine if metapopulation theory is applicable; such frameworks for other large mammals in habitat patches have been developed (e.g., Elmhagen and Angerbjérn 2001). Occupancy by Gray Wolves on islands likely is influenced by the presence and abundance of deer, and security from exploitation by humans. These fac- tors might be mediated by size of island, isolation, topography, shape, and extent of logging. Because wolves are obligate predators of ungulates, islands without deer would not sustain wolves. Persistence times, however, are probably a function of the abun- dance and availability of deer (Fuller 1989) as well as island size and isolation. Wolves seem unable to per- sist indefinitely on small and isolated islands, even if deer are present. On isolated islands, they may be subject to a “minimum area effect” (sensu Lomolino 1986). That is, although their immigration potential is high, they can only maintain populations on the largest of isolated islands. In the 1960s, biologists introduced four Gray Wolves to the 73 km? Coronation Island, southeast Alaska, 900 m from another landmass, to study the effects on resident deer (Klein 1996). After reaching a peak of 13 animals in four years, the wolf popula- tion, having severely reduced deer numbers, plum- meted to one. Klein (1996) and Person et al. (1996*) stated that small and isolated islands, such as Coronation, are unable to support populations of wolves. Person (2000) noted that islands as large as 180 km? with deer carrying capacities greater than 2500 in southeast Alaska did not continuously sup- port wolves between 1955 and 2000. Lamolino (1986) argued that extinction-prone species, such as large carnivores, might be common on small islands only if their immigration rates are high relative to extinction rates. This can apply to Gray Wolf packs as a conceptual unit. The social group may be sustained if their home range includes a collection of nearby islands (Klein 1996; Person et al. 1996*; Person 2000). Including multiple landmasses would compensate for extirpation risk on individual islands. Of the four most isolated islands on which we noted sign, wolves currently were present only on Dundas and Dunira Islands (Figure 1; Table 1), and probably formed one social group. Although both are isolated from large landmasses, collectively these and smaller nearby islands total 239 km2, an area within the range of home ranges calculated for wolves of southeast Alaska (Person 2000). The viability of wolves on large but isolated islands » or island groups such as Dundas and Dunira may be uncertain. For example, although the 540 km? Isle Royale has supported wolves since the late 1940s THE CANADIAN FIELD-NATURALIST Vol. 116 ve without further immigration, the population has expe- rienced considerable declines due to disease, demo- graphic stochasticity, and shortage of food (Peterson and Page 1988; Vucetich et al. 1996; Peterson et al. 1998). Moreover, continued viability may be threat- ened by a loss of genetic diversity (Wayne et al. 1991). In coastal British Columbia, an island’s topography would interact with size and isolation in determining the abundance of deer and thus the viability of wolf populations. The complex topography of many inner coastal islands can include elevations of 1100-m or more, above which deer in adjacent southeast Alaska are not known to occur (Schoen and Kirchhoff 1985). Thus, habitat suitable for deer and wolves can be less than predicted based on area alone. Mounting evidence suggests that carrying capacity for deer will be reduced by even age stand manage- ment in the Pacific Northwest because clearcut log- ging is largely incompatible with the food require- ments of deer (Wallmo and Schoen 1980; Alaback 1982; Rose 1982; Alaback 1984*; Schoen et al. 1984, 1998; Van Horne et al. 1988; Hanley et al. 1989*). Most notably, closed canopies that develop 20-30 years after harvest severely limit forage for deer and persist for 140-160 years (Wallmo and Schoen 1980; Alaback 1982; Schoen et al. 1988). Persistence on islands might also be influenced by human-caused mortality. Island wolf populations are vulnerable to over-exploitation because water bodies may slow immigration and re-colonization (Person et al. 1996*; Person 2000). Person (2000) reported that humans on Hecata Island, southeast Alaska, nearly exterminated Gray Wolves on two occasions. In our nearly roadless study area, the shape of islands may affect human-caused mortality. Coastlines are analo- gous to roads because travel routes of human hunters in boats and wolves converge. Because of high coastline to interior ratios, long and narrow islands provide less security than rounder islands. Although annual human-caused mortality in the study area is only about 2 %, the human population is growing considerably and hunting regulations are liberal (Darimont and Paquet 2000*). Moreover, more road building is planned to support industrial logging (Darimont and Paquet 2000*). This is a con- cern, aS many authors have shown the lethal impact of roads on Wolves (i.e., Thiel 1985; Jensen et al. 1986; Fuller 1989; Paquet et al. 1996*; Callaghan 2002) and other wildlife species (review in Trom- bulak and Frissell 2000). We encourage others to invoke a broad spatial and temporal perspective when considering the distribu- tion and conservation status of this population of Gray Wolves, regardless of their wide distribution in the study area now. Extant populations, such as those in coastal British Columbia, form remnant groups of those that first colonized North America roughly 700 000 years ago (Nowak 1979; Kurten and Ander- son 1980) and historically ranged in every habitat that 2002 DARIMONT AND PAQUET: WOLVES OF BRITISH COLUMBIA’S COAST 421 supported ungulate prey. During the last few hundred years, humans have reduced Gray Wolf populations and effectively isolated them to remote areas of Canada, Minnesota, and Alaska (Mech 1970, 1995). On the West Coast, wolves were extirpated in Cali- fornia, Oregon, and Washington, as well as in the Lower Mainland region of British Columbia (Theberge 1991). Wolves of these coastal rainforests are now restricted to southeast Alaska and most of British Columbia. Furthermore, coastal biota, includ- ing the Gray Wolf, is thought to be at least partially isolated from continental populations by coastal mountain ranges (e.g., McCabe and Cowan 1945; Klein 1965; MacDonald and Cook 1996; Person et al. 1996*; Byun et al. 1997; Conroy et al. 1999; Dari- mont and Paquet 2000*; Cook and MacDonald 2001; Figure 1). Finally, wolves of coastal British Columbia should be considered a natural source population for those in nearby southeast Alaska, a population for which there is considerable conservation concern due to timber removal and associated effects (Kirchhoff 1991*; Person and Ingle 1995*; Person et al. 1996*; Person 2000) Acknowledgments This study took place in the Traditional Territories of several First Nation groups, from whom we sought permission before research began. We are grateful to the Raincoast Conservation Society for financial and logistical support, as well as for their invaluable local knowledge. We are obliged to Gudrun Pfleuger, Chester Starr, and numerous others for fieldwork, and to Rob Boethe, Heather Bryant, and Sabrina Lundquist for island area and isolation estimates. While preparing the manuscript, CTD was supported by a Natural Sciences and Engineering Research Council (NSERC) — Industrial Post- graduate Scholarship. 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Hansen, R. O. Peterson, U. S. Seal, A. Eisenhawer, L. D. Mech, and R. D. Krumenaker. 1991. Conservation genetics of the endangered Isle Royale gray wolf. Conservation Biology 5: 41-51. Weaver, J. L., and S. H. Fritts. 1979. Comparison of coy- ote and wolf scat diameters. Journal of Wildlife Manage- ment 3: 786-788. Williamson, M. H. 1981. Island Populations. Oxford Uni- versity Press. New York, NewYork. Received 4 March 2001 Accepted 23 June 2002 A Review of the Status of the Atlantic Wolffish, Anarhichas lupus, in Canada* NIALL R. O’ DEA and RICHARD L. HAEDRICH! Memorial University of Newfoundland, Department of Biology, 4 Clark Place, St. John’s, Newfoundland AlC 5S7 Canada 'Corresponding author O’Dea, Niall R., and Richard L. Haedrich. 2002. A review of the status of the Atlantic Wolffish, Anarhichas lupus, in Canada. Canadian Field-Naturalist, 116(3): 423-432. The Atlantic Wolffish, Anarhichas lupus Linneaus, 1758, is a large blenny-like marine fish found in moderately deep and cold waters of the North Atlantic on rocky and hard bottoms from the Gulf of Maine to northwestern France. Wolffish was once the target of a directed fishery, but is not at present; nonetheless, it continues to be taken incidentally as by-catch in the trawl fishery. In Canadian waters, it has been most abundant off Newfoundland and Labrador, where it is a characteris- tic member of the deep cold-water fish assemblage on the continental shelf, but its numbers there, as indicated by scientific surveys, declined by 91% between 1978 and 1994. In other Canadian areas, where wolffish abundance has historically been lower by about an order of magnitude, populations appear stable. Wolffish are relatively sedentary and slow-growing. They make nests, and guard their large eggs. They feed mostly on bottom invertebrates. Life histories and distribution pat- terns can vary considerably across the species’s range. Aggressive trawl fisheries, now in abeyance by the imposition of widespread moritoriums, appear to have had an impact on wolffish numbers. In the Newfoundland region, numbers have declined steadily in scientific surveys, the number of locations where the species occurs is fewer, the range where abundant appears to be shrinking, and the mean size has decreased so that mature individuals have become relatively rare. Slow growth, fidelity and inshore nesting habit, and limited dispersal make rescue unlikely, and bottom trawling and dredging have probably damaged habitat. Key Words: Atlantic Wolffish, Anarhichas lupus, North Atlantic, population size, status, Newfoundland, Labrador. The wolffishes, family Anarhichadidae, are large blenny-like marine fishes which inhabit moderately deep waters of the North Atlantic and North Pacific oceans. They are named for the characteristic large, conical, canine-like teeth employed in consuming the benthic crustaceans and invertebrates which are their chief food source. Three species of wolffish occur in the Canadian Atlantic area: the Northern Wolffish (Anarhichas denticulatus Kr@yer, 1844), the Spotted Wolffish (Anarhichas minor Olafsen, 1774), and the Atlantic Wolffish (Anarhichas lupus Linnaeus, 1758). The Atlantic Wolffish is an elongate, laterally compressed fish with a heavy head, blunt snout and rounded profile. It can reach a length of 150 cm and a weight of almost 20 kg. In common with all wolffish, it has distinctive and prominent large canine-like teeth in the front of the jaws and flat- tened, grinding teeth behind. There are no pelvic fins. Atlantic Wolffish vary in colour, depending on their surroundings, from slaty blue to dull olive green to purplish brown, and there are dark trans- verse bars on the body (Whitehead et al. 1986; Scott and Scott 1988). The Atlantic Wolffish can be distinguished from the other two wolffishes of the region by the nine to thirteen irregular and broken dark transverse bars on its body, some of which extend onto the dorsal fin. As well, its musculature is firm, not jelly-like as in the Northern Wolffish, and the grinding teeth on the vomer extend to the rear of the mouth behind the rows of palatine teeth (Barsukov in Whitehead et al. 1986). Distribution Adapted to a broad range of depth and tempera- ture, the Atlantic Wolffish is widely distributed on both sides of the North Atlantic. In the eastern North Atlantic it is found from Iceland, the Faeroes, Spitzbergen, the White Sea, and Murman coast, south to the British Isles and the western coast of France (Wheeler 1969; Whitehead et al. 1986). In the western North Atlantic, it is found off west Greenland and southern Labrador, in the Strait of Belle Isle and the Gulf of St. Lawrence, off the east and west coasts of Newfoundland and on the Grand Banks. Its most southerly range extends from the Scotian Shelf to the Gulf of Maine with occasional strays off New Jersey (Bigelow and Schroeder 1953; Scott and Scott 1988). Thus the Atlantic Wolffish is basically a coldwater fish. Mahon et al. (1998) have identified it as a characteristic member of the “northern, cold, deep, aggregated” demersal fish assemblage that occurs on the continental shelves off northeastern Newfoundland and southern Labrador. The East Coast of North America *The status report on which this paper is based was reviewed and approved by COSEWIC in November 2000; status assigned — Special Concern. 423 424 Strategic Assessment Project (ECNASAP) on-line Groundfish Atlas (http://www-orca.nos. noaa.gov/ projects/ecnasap/ecnasap.html) summarizes twenty years of distributional data from scientific research surveys conducted in the western North Atlantic in the map ATLWOL.GIF. The map shows the distri- bution of Atlantic Wolffish in the western Atlantic is predominately Canadian. Within that area, it is most prevalent and abundant on the deep shelf off north- eastern Newfoundland and Labrador. Protection Because the Atlantic Wolffish is not at present the target of a directed fishery in the western Atlantic it is unmanaged and there are no specific mechanisms, such as total allowable catch limits, in place that afford it protection. However, the Canadian Atlantic groundfish moratorium imposed in 1992 in response to the collapse of Atlantic Cod, Gadus morhua, may have provided some temporary indirect protection for the wolffish by reducing overall trawling pressure. The Atlantic Wolffish was listed (November 2000) as “Special Concern” by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC), but not by The World Conservation Union (IUCN), or the Convention on International Trade in Endangered Species (CITES). The biology of the Atlantic Wolffish may provide it with some limited protection. As a benthic organ- ism which feeds on the bottom, it is unlikely to be caught in trawls above the ocean floor. Additionally, it frequents caves and crevices between and under large rocks, potentially affording at least some protection from bottom trawls and dredges. Population Size and Trends The most important data for assessing the status of Canadian marine fishes under COSEWIC are those which document numerical decline. There is a wealth of such data, which come from regular scientific surveys conducted by government and are expressly designed to monitor changes in abundance of demersal fishes. The same data can also be used to generate secondary but also useful information on possible changes in the number of fish per trawl, the mean size of the fish and the range and habitat. Data from random-stratified scientific survey trawls off eastern Newfoundland (Atkinson 1994) were provided by Canada’s Department of Fisheries and Oceans (DFO) and summarized by Villagarcia (1995; also Haedrich and Barnes 1997). The surveys are intended mainly to assess the size of commercial fish stocks, but they also catch most species in the demersal fish community (Brown et al. 1996). The number of individual trawls made (the number of stations) in any one year can be over a thousand. Subsequently, in July to September 2000, DFO made large amounts of trawl survey data and analyses THE CANADIAN FIELD-NATURALIST Vol. 116 - available from all Canadian areas specifically for our Atlantic Wolffish assessment. The number/tow (which fishery biologists refer to as the “catch per unit effort” or CPUE) from scientif- ic surveys is used as an index of population size. Over the period from 1978 to 1993, this index was calculated as the total number of Atlantic Wolffish caught off Newfoundland in a year divided by the total number of stations sampled at appropriate depth and temperature ranges for the species in that year. The appropriate depth and temperature ranges for the Atlantic Wolffish are determined using the niche axis approach developed by Fischer and Haedrich (2000) and represent the ranges of those two envi- ronmental parameters within which the wolffish is most likely to be encountered; i.e., 100 to 400 m for depth and greater than or equal to -0.5°C for tem- perature. The scientific survey data from Newfoundland indicate a sharp decline in the Atlantic Wolffish population. In 1978, the number/tow averaged 10.5 individuals caught in each tow. The following year it dropped more than 25% to 7.1 individuals per tow. It then continued to decline steadily, and by 1993 had fallen to just 0.96 individuals per tow. Over the full period from 1978 through 1993, 16 years or about two wolffish generations, the number/tow declined by 91% (Figure 1). Kulka and DeBlois (1997) note the same general decline for all Newfoundland statistical areas in the period from 1987 to 1995. Since 1995, scientific survey data that include information on Atlantic Wolffish continue to be gathered by DFO. For the Scotian Shelf and Gulf of St. Lawrence, recent catch rates are comparable to ones observed earlier and are low, on average < 1.0 individuals per tow. In the northern Gulf, Atlantic Wolffish are also not very abundant (Powles 2000, personal communication) the average catch rate for 1990-1999 is 0.42 fish per tow. The abundance of Atlantic Wolffish in these more western regions has been low throughout the whole of the past 20 years, and has not shown the decline apparent off Newfoundland where the wolf-fish was once much more abundant than it is today (Figure 1) and where, as the ECNASAP map shows and Mahon et al. (1998) indicate, this species assumes its greatest importance in the Northwest Atlantic. DFO’s main population assessment tool is a Stratified Random Assessment Program (STRAP). This computer analysis takes catches from at least two trawls within defined strata, scales them accord- ing to the total area of the stratum (within which the species is assumed to be uniformly abundant), and calculates an estimated number of fish presumed present. To get a total, those numbers are summed across all strata where the fish was encountered. The size of a single stratum can range from 30 to 2817 square nautical miles (average = 697) and, because a single survey trawl sample covers about 0.009 2002 O’ DEA AND HAEDRICH: ATLANTIC WOLFFISH IN CANADA 425 Number/Tow VMMMMMMlllld ld ANA Y RG teBOw B200 284: B69) 88 90 = 92 94 NSS WN NING 96 SR. UU Year Atlantic wolffish, mean CPUE +/- std err Number/Tow 78 79 8 81 8& 8 8&4 8 8 87 8 8 90 91 92 Year FicurE 1. Top — Bars: CPUE (no./stn in each year) for Atlantic Wolffish, Anarhichas lupus, caught in tows within appropriate depth and temperature ranges, 1978-93; Line, X’s — same, all stations (data provid- ed by DFO in September 2000). Bottom — mean CPUE + standard error for all positive tows. Eastern Newfoundland, 1978-1993, DFO Fall Survey Data. Square nautical miles and there are on average two trawls per stratum, the scale-up is prodigious (Schneider et al. 1999). The summed estimates from STRAP analysis results for Atlantic Wolffish in all Canadian waters across the period 1978 to 1999 are shown in Figure 2. Because the sampling protocol changed in 1995, values from Newfoundland areas after 1994 are divided by a correction factor for comparability. For demersal species like wolffish, that factor ranges in general from 3.1 for adults to 10.7 for juveniles (Bundy et al. 2000). Calculated for the Atlantic Wolffish alone using Bundy et al.’s (2000) formula, the factor is 4.85. The STRAP results also indicate a large decline in the wolffish population, down 87% over the 16 years from 1978 to 1994 (about two wolffish generations, and with no change in sampling protocol). From 1978 to 1999 (a little less than three wolffish generations 1995-1999 Newfoundland catches adjusted by 4.85), the decline indicated is 83%. Despite the debatable value of STRAP’s absolute numbers, the annual estimates 426 Millions of Wolffish 78 80 82 84 86 88 90 92 94 96 98 00 Year FIGURE 2. Twenty years of STRAP analyses for Atlantic Wolffish, Canadian waters. Solid line, filled circles: Engels trawl estimates. From 1978 to 1994 (16 years; i.e., about two wolffish generations) there was no change in sampling method, and the decline is 87%. Dotted line, small dots: corrected estimates from Campelen trawls using the factor’ 4.85 (see text), 1995-1999. Based on information provided by DFO in September 2000. Straight line shows the COSEWIC endangered criterion. % Frequency 100 80 60 40 20 <15 THE CANADIAN FIELD-NATURALIST Vol. 116 - turn out to be well-correlated (r = 0.93) with the sim- ple metric we prefer, the number/tow. From 1978 tto 1993, the relative frequency of high survey catch rates off Newfoundland has declined, and low catch rates have been on the rise (Figure 3). In 1978, catches of five or less wolffish in a tow constituted less than 40% of all catches; by 1984 the frequency of low catches (<5) had risen to more than 70%, and in 1993 it was almost 90%. The increased frequency of low catch rates is another indication of declining population density. The trend to a declining mean size of fish, consid- ered together with the declining numbers, is yet another indication that wolffish populations are in trouble. These trends vary according to area, but all are down. The average size of Atlantic Wolffish from the Scotian Shelf and southern Gulf of St. Lawrence, while showing inter-annual variation, declined overall by 50% or more from the mid- 1980s to the present; the trend lines are essentially the same. Off Newfoundland, mean wolffish size declined from around a kilo in 1978 to near 700 gm in 1993. Fish of that size are probably not yet 1978 1984 1993 <20 <25 <30 >=30 Catch Rate Class (number/tow) FIGURE 3. Percent frequency of large and small catches of Atlantic Wolffish, Anarhichas lupus, by catch rate classes of 5, off eastern Newfoundland, 1978-1993. DFO Newfoundland Fall Survey Data. 2002 O’ DEA AND HAEDRICH: ATLANTIC WOLFFISH IN CANADA 427 mature. In the Northern Gulf, the average size in the period 1990-1999 is less than 500 gm. To the north in Greenland waters, scientific trawl surveys (Ratz 1997; Moller and Ratz 1999) have been done differently than in the U.S. and Canada, so CPUEs are not directly comparable. Biomass and abundance of Atlantic Wolffish off East Greenland increased somewhat from 1982 to 1998, but off West Greenland (the region nearest to Canada), biomass has declined by about an order of magnitude and numbers are down by about half. Mean age in the whole Greenland population in 1998 (= 4 yr) is half of what it was in 1982 (Moller and Ratz 1999), so the picture appears to be quite similar to that seen off Newfoundland. To the south, Atlantic Wolffish have never been very abundant in U.S. waters (CPUE <0.2 individuals per tow). There, wolffish did expe- rience a significant decline in the mid-1970s, several years before one of much greater magnitude in Atlantic Canada. The period since 1992 is an anomalous one for all Canadian waters. Relative to the past, fish popula- tions are at an all-time low. For that reason, bans on fishing (moratoriums) have been in effect in most regions for various periods of time, and these contin- ue. Fishing predation is thus much relaxed, and pop- ulations should do better as long as that situation continues, which will not be forever. A cornerstone in the Fisheries Resource Conservation Council’s (FRCC) approach to management (this quasi-inde- pendent group advises the Minister on the status of commercial fish stocks) is adherence to the precau- tionary principle “when in doubt, err on the side of the fish” (FRCC 1996). 70 2 60 c= e) s 50 -- boo net ste ee sce s speebetescte 28 see apt eine gel ie ap rae arte a = = 40 Nn = ‘ S30 NE deg Ge oe) B20 fn ge vo 5 2 10 0 iba. 80), 82 84 86 88 90 92 94 96 98 Year FIGURE 4. Percentage of scientific survey stations that captured Atlantic Wolffish, Anarhichas lupus, 1978-1999. Solid line, circles: Canadian waters, ECNASAP data. Dotted line, Xs: Newfoundland waters only, data provided by DFO in September 2000. Note: sampling protocol changed in 1995. General Biology Habitat The Atlantic Wolffish is found principally in the deep waters of the continental shelf on rocky or hard clay bottoms, and only occasionally on sand or mud. Like other wolffish species, its migrations are local and limited (Templeman 1984) and it does not form large schools. It is, however, known to perform small seasonal inshore-offshore migrations (Keats et al. 1985). It is found anywhere from very shallow water to 500 m deep and is said to prefer depths between 100 and 150 m; this apparent preference, however, varies depending on the locality. The Atlantic Wolffish is a cool to cold water fish, tolerating a broad temperature range of -1°C to 10°C, though pre- ferring temperatures between -0.4°C and 6°C. These preferred temperature ranges vary with locality. The percentage of all annual DFO survey stations where the Atlantic Wolffish was actually caught declined steadily from near 35% in 1978 to about 10% in 1994 (Figure 4). For the area off Newfoundland, the Atlantic Wolffish occurred in 88% of the stations where it was expected (according to preferred depth and temperature ranges) in 1978, and that level of occurrence continued until about 1985. After 1985, the percentage declined steadily to only 33% in 1993. Reproduction Spawning in the Atlantic Wolffish varies in time and place. Off the east coast of Newfoundland some Atlantic Wolffish migrate to shallow inshore waters in spring, spawning in September with hatching occur- ring by mid-December (Keats et al. 1985). In both the Gulf of Maine (Nelson and Ross 1992) and eastern Newfoundland (Keats et al. 1986), it is thought that only fish over 50 cm and sexually mature move inshore, while smaller, juvenile fish remain in deep water. In Greenland the maximum number of spawn- ers were observed by Beese and Kandler (1969) in September and October. In Iceland the situation is quite different from that in Newfoundland; Jonsson (1982) reports that the Atlantic Wolffish moves from shallow into deeper waters to spawn from September to December or January, subsequently returning to shallow waters to feed. Similarly, spawning and feed- ing are exclusive events in the White Sea, with spawning occurring in the deep waters between 70 m and 300 m from August through September (Pavlov and Novikov 1993). Keats et al. (1985: 2567) point out that the literature “suggests geographical and depth-related variability in the reproductive season of Atlantic Wolffish”. As such, discrete geographical populations of Atlantic Wolffish may have distinct and different life histories, a possibility that requires further study. Wolffish lay some of the largest eggs of any fish known — up to 6.0 mm in diameter. The eggs are deposited in a large mass on the bottom and are 428 guarded by the male. Larvae remain mostly close to the bottom, rarely swimming to the surface and tend- ing to remain close to the site of hatching. Limited adult migrations and the restricted disper- sal of larvae from their hatching site are potential risk factors for the survival of Atlantic Wolffish populations on the smaller scale. If the population of a given region is decimated through environmental or anthropogenic causes, it is unlikely to be replen- ished by populations from elsewhere. The broad range of temperature and depth to which the Atlantic Wolffish appears to be adapted may be indicative of the unique depth and temperature regimes of discrete and separate, rather than confluent, populations. Growth Growth rates of the Atlantic Wolffish in Can- adian Atlantic waters are unknown and there are lit- tle data available for other areas. In Europe, it takes a fish three years to reach 25 cm TL (Wheeler 1969), and Scott and Scott (1988) report that growth rates in Atlantic Canada are slow after the first’ summer. Growth slows even further at five to six years of age when energy is diverted to gonadal development (Nelson and Ross 1992). Off eastern Newfoundland the Atlantic Wolffish reaches maturity between 43 and 67 cm TL (weights of 0.56 and 2.39 kg); fifty percent of individuals between 52 and 60 cm TL (1.02 and 1.57 kg) are mature (Templeman 1986). Fish of this size are 8 to 10 years old. Atlantic Wolffish are known to reach 152 cm in length and specimens aged to twenty years have been taken off Iceland (Scott and Scott 1988). Feeding The Atlantic Wolffish feeds primarily on hard- shelled benthic invertebrates such as echinoderms, molluscs, and crustaceans. It is thought to be a key player in the ecosystem, known to control the densi- ty and spatial distribution of species such as Green Sea Urchins, Strongylocentrotus droebachiensis (Hagen and Mann 1992), crabs (Witman and Sebens 1992) and Giant Scallops, Placopecten magellanicus (Stokesbury and Himmelmann 1995). The Atlantic Wolffish also consumes small amounts of fish, particularly Redfish, Sebastes sp. Young wolffish eat echinoderms almost exclusively but this food source becomes less important as the fish grows. While mature males reduce feeding close to spawning time and until their egg-guarding duties have ceased, females reduce feeding as gonads mature, but resume feeding immediately following spawning (Keats et al. 1985). Little is known about what preys on the Atlantic Wolffish, but juvenile specimens have been reported in the stomach contents of cod (Scott and Scott 1988). Limiting Factors There are no direct studies of factors responsible for the declines observed in wolffish abundance. THE CANADIAN FIELD-NATURALIST Vol. 116 rs Following the Northern Cod collapse off Newfoundland in 1992, a number of causes for the cod decline were suggested, including especially environmental changes. On a community level, simi- lar declines in both commercial and non-commercial fish species there seemed difficult to understand without invoking such causes (Gomes et al. 1995). The consensus now is that environment may have played some role, but overfishing was clearly the major cause of the declines observed in most groundfish species (Sinclair and Murawski 1997; Villagarcia et al. 1999). When assessed over only a slightly longer time scale, fishing in the area is argued to have been responsible for the extraordi- nary decline of the large and once abundant and widely-distributed Barndoor Skate, Raja laevis (Casey and Myers 1998). The Atlantic Wolffish has been landed from com- mercial fisheries for many years. Once a directed tar- get of the western Greenland fishery, it is now taken incidentally as by-catch only. There, M6ller and Ratz (1999) report that calculated wolffish mortality is positively correlated with commercial landings of cod and shrimp. This indicates that fishing for other species can have a negative impact on wolffish through removal in the by-catch. There is no directed fishery for the Atlantic Wolffish in Canadian waters, though it is taken as by-catch by offshore trawlers. Recently, wolffish filets have begun to appear in St. John’s fish shops, and there is concern that with developing markets “fishermen might target wolffish as a ‘directed by-catch’”(FRCC 2002: 46). In fisheries data compiled by the Food and Agriculture Organization of the United Nations (FAO), wolffish landings for the western North Atlantic are reported for the whole family (three species) rather than for each individual species; the Atlantic Wolffish comprises the majority of those landings in Canada (Kulka and DeBlois 1997). While the northeastern Atlantic fishery (apparently focussed more on the Spotted Wolffish, A. minor; Wheeler 1969) has had annual landings of around | 30 000 tonnes since the late 1950s, with two peaks | of over 50 000 tonnes in 1962 and 1974, the north- western Atlantic fishery has always been smaller. Northwest Atlantic landings (Figure 5) hovered around 5000 tonnes through the 1950s and then rose through the 1960s and 1970s to a peak of 22 000 tonnes in 1979. Landings then declined steadily through the 1980s and 1990s; in 1984 they stood at 6000 tonnes and by 1996 had fallen to 1700 tonnes. The principal countries involved throughout in the history of the wolffish fishery have been Canada and Greenland, each with about a third of the landings from 1950 through 1996. The Soviet Union and East Germany also played significant roles in this fishery through the 1960s and 1970s, with East Germany responsible for almost 70% of 1979’s record catch. 2002 O’ DEA AND HAEDRICH: ATLANTIC WOLFFISH IN CANADA 429 Landings, 1000 t 25 20 15 [Portugal E=jEastern Bloc }- WAGreenland Canada £3 Other ee | Y Year FiGuRE 5. History of the northwest Atlantic fishery for Atlantic Wolffish, Anarhichas lupus, 1950-1996. FAO Data. The Soviet Union and East Germany were essentially out of the picture by the early 1980s, and by 1990 Greenland had seriously curtailed its fishing efforts as well. In recent years, from 1990 through 1996, Portugal has become the major participant in a much reduced fishery. Apart from the direct adverse impact of fisheries on Atlantic Wolffish, human activities also have indirect and detrimental effects on this species. The groundfish trawls, generally otter trawls, in which wolffish are caught can result in incidental mortality and damage to fish which come in contact with the mobile fishing gear but are not caught. Perhaps even more importantly, the steel doors or otterboards of the net, along with heavy bottom lines and rollers, scour the seabed as they are dragged across it (Watling and Norse 1998). This practice may cause significant habitat damage by removing or re- distributing the rocks and boulders under which these fish shelter, spawn and build nests. Studies on Georges Bank (Collie et al. 1997) and in the Gulf of Maine (Auster et al. 1996), areas within the southern limit of the Atlantic Wolffish’s range, show the considerable degree of damage that can result from bottom dragging there. Jennings and Kaiser (1998) provide an excellent overview of the entire question of fishing impacts on habitat; they point out that these can vary quite a bit depending on local condi- tions, but suggest that the greatest and most lasting impacts are most likely to occur on hard substrates in deep water; i.e., just those habitats favoured by th Atlantic Wolffish. i Bottom trawling for fish and dredging for scallops and clams, in addition to digging up and disrupting bottom habitats, also re-suspends bottom sediments, which can smother spawning areas and damage gills. Re-suspension of sediments may change the sedi- ment chemistry or release settled toxic heavy metals. Other human activities such as channel dredging and aggregate extraction may do considerable harm to bottom habitats by destabilizing the seabed, increas- ing erosion, and polluting previously healthy areas (Messieh et al. 1991). Status Determination Under IUCN Categories and Criteria, criterion A: “Declining Population”, the Atlantic Wolffish, with its 87% decline rate over a bit more than two generations, would fall in the category Critically Endangered, defined as a “population decline rate at least 80% in 10 years or three generations” (Hilton-Taylor 2000: 55). Musick (1999) feels that marine fish, by virtue of their widespread distribution and thus relatively great abundance, should be treated differently from other species in respect to consideration as species at risk. He proposes an approach that involves two steps, first determining the productivity class of a species (based on growth, fecundity and age charac- teristics) and then classifying it on the basis of arbitrary decline thresholds. The Atlantic Wolffish, on the basis of age at maturity and lifespan, falls in Musick’s (1999) “Low” productivity category, and exhibits a decline of 87% over less than three gener- ations, just over the 85% decline threshold for 430 species in the Low category. According to Musick’s proposed scheme, the Atlantic Wolffish would be listed as Vulnerable and would then be subjected to closer scrutiny for final classification. That scrutiny would explicitly consider issues concerning the possibility of a shrinking range, local distribution, nesting habit, and possibility of habitat destruction. The data available on all these issues arguably support upgrading. Hutchings (2000) shows that exemption of marine fish from established species-at-risk criteria cannot be supported on the basis of empirical data and, furthermore, to do so would be inconsistent with a precautionary approach to fisheries management and the conservation of biodiversity. He further argues that extinction risk alone is not very useful from either a management or an ecological perspective and that the other face of the issue, the probability of recovery, should be weighed seriously in assigning status. He suggests a classification scheme that would replace Vulnerable to Critically Endangered with Conservation Categories ranging from Priority I to IV. The empirical data show that population recovery in marine fish is negatively correlated to the magnitude of population decline. With its high decline rate of 87%, the likelihood that Atlantic Wolffish populations would show signs of recovery after 15 years is very low, and this species would fall in Hutching’s Conservation Category Priority IV, his highest rating. The FRCC and the North Atlantic Fishery Organisation (NAFO) have found Atlantic Wolffish populations to be in poor shape. In the far north of Canadian Atlantic waters (subarea 1), NAFO (2000) notes that Atlantic Wolffish remains severely deplet- ed and recommends that there be no fishery for the species in 2001 and 2002. FRCC (2000) indicate that for Atlantic Wolffish on the Scotian Shelf the stock is low, the biomass is declining, growth and condi- tion are below average and the age structure is poor. They recommend that there be no fishery for the next two years and that restrictive measures be put in place to minimize by-catch, noting that “concentrat- ed fishing effort . . . has likely contributed to overall decline” (FRCC 2000: 42). The DFO scientific survey data show an 87% decline in the number of Atlantic Wolffish in Canadian waters over the 16-year period (about two wolffish generations) from 1978 to 1994. Under cri- terion A: “Declining Population” of COSEWIC’s Risk Categories and Criteria, this population decline rate places the Atlantic Wolffish clearly in the cate- gory Endangered, defined as a “population decline rate at least 50% in 10 yrs or 3 generations” (Haedrich, 2001: appendix page 2). At its annual meeting in November 2000, COSEWIC considered the Atlantic Wolffish, and after an extended debate voted to list it as Special Concern. The statement accompanying that designa- RE Se eee ae oe bt 5 THE CANADIAN FIELD-NATURALIST Vol. 116 tion read: “The total population of this large, solitary, slow-growing, late-maturing, nest-building benthic fish has declined significantly since the 1970’s. Apparent threats are related to fishing and habitat alteration, perhaps compounded by environ- mental change. Met criterion for Endangered Alb, but because it is not considered to be facing immi- nent extinction, it was listed as Special Concern on the grounds that it is a species that would likely become at risk if not for active protection and management.” Acknowledgments Our research was supported in part by an under- graduate research award and an operating grant from the Natural Science and Engineering Research Council of Canada (NSERC). We thank David Kulka, DFO St. John’s, for help in obtaining and understanding recent assessment data, and Ivone Figueiredo, Instituto de Investagagao das Pescas e do Mar (IPIMAR) Lisbon, for discussions on the statis- tical ins and outs of fisheries analyses. Kelly Barrington helped immensely with final data reduc- tion. The data and supporting materials used in the preparation of this report have been deposited in the Newfoundland Archive, Memorial University of Newfoundland. Literature Cited Atkinson, D. B. 1994. Some observations on the biomass and abundance of fish captured during stratified random bottom trawl surveys in NAFO divisions 2J and 3KL, Autumn 1981-1991. NAFO Scientific Council Studies 21: 43-66. Auster, P. J., R. J. Malatesta, R. W. Langton, L. Watling, P. C. Valentine, C. L. Donaldson, E. W. Langton, A. N. Shepard, and I. G. Babb. 1996. The impacts of mobile fishing gear on seafloor habitats in the Gulf of Maine (Northwest Atlantic): implications for conservation of fish populations. Reviews in Fisheries Science 4: 185-202. Beese, G., and R. Kandler. 1969. Beitraége zur Biologie der drei nordatlantischen Katfischarten Anarhichas lupus L., A. minor Olafs. und A. denticulatus. Berichte Deutscher Wissenschaftliche Kommission fuer Meeresforschung 20: 21-29. Bigelow, H. B., and W. C. Schroeder. 1953. Fishes of the Gulf of Maine. United States Government Printing Office, Washington DC, 577 pages. Brown, S. K., R. Mahon, K. C. T. Zwanenberg, — D. B. Atkinson, K. R. Buja, L. 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Carscadden, and J. E. Simon. 1996. Recent excursions of capelin (Mallotus villosus) on the Scotian Shelf and Flemish Cap during anomalous hydro- graphic conditions. Canadian Journal of Fisheries and Aquatic Sciences 53: 1473-1486. FRCC (Fisheries Resource Conservation Council). 1996. Building the Bridge: 1997 Conservation Requirements for Atlantic Groundfish. Report to the Minister. Department of Fisheries and Oceans, Ottawa. FRCC (Fisheries Resource Conservation Council). 2000. 2000 Conservation Requirements for Groundfish Stocks on the Scotian Shelf and in the Bay of Fundy (4vwx), in Sub-areas 0, 2 + 3 and Redfish Stocks. FRCC.2000.R.1. Report to the Minister. Department of Fisheries and Oceans, Ottawa. FRCC (Fisheries Resource Conservation Council). 2002. 2002/2003 Conservation Requirements for Groundfish Stocks on the Scotian Shelf and in the Bay of Fundy (4vwx), in Sub-areas 0, 2 + 3 and Redfish Stocks. FRCC.2002.R.1. Report to the Minister. 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T., and K. H. Mann. 1992. Functional response of the predators American lobster Homarus americanus (Milne-Edwards) and Atlantic Wolffish Anarhichas lupus (L.) to increasing numbers of the green sea urchin Strongylocentrotus droebachiensis. Journal of Experimental Marine Biology and Ecology 159(1): 89-112. Hilton-Taylor, C. Compiler. 2000. 2000 IUCN Red List of Threatened Species. IUCN, Gland, Switzerland and Cambridge, U.K. xviii + 61 pages. Hutchings, J. A. 2000. Collapse and recovery of marine fishes. Nature 406: 882-885. O’ DEA AND HAEDRICH: ATLANTIC WOLFFISH IN CANADA 431 Jennings, S., and M. J. Kaiser. 1998. The effects of fishing on marine ecosystems. Advances in Marine Biology 34: 201-352. Jonsson, G. 1982. Contribution to the biology of catfish (Anarhichas lupus) at Iceland. Rit Fiskideildar 6: 2—26. Keats, D. W., G. R. South, and D. H. Steele. 1985. Reproduction and egg guarding by Atlantic Wolffish (Anarhichas lupus: Anarhichadidae) and ocean pout (Macrozoarces americanus: Zoarcidae) in New- foundland (Canada) waters. Canadian Journal of Zoology 63: 2565-2568. Keats, D. W., G. R. South, and D. H. Steele. 1986. Where do juvenile Atlantic Wolffish, Anarhichas lupus, live? Canadian Field-Naturalist 100: 556-558. Kulka, D. W., and E. M. DeBlois. 1997. Non-traditional groundfish species on Labrador Shelf and Grand Banks: Wolffish, Monkfish, White Hake, and Winter (Blackback) Flounder. DFO Atlantic Fisheries Research Document 96/97. 49 pages. Mahon, R., S. K. Brown, K. C. T. Zwanenburg, D. B. Atkinson, K. R. Buja, L. Claflin, G. D. Howell, M. E. Monaco, R. N. O’Boyle, and M. Sinclair. 1998. Assemblages and biogeography of demersal fishes of the east coast of North America. Canadian Journal of Fish- eries and Aquatic Sciences 55: 1704-1738. Messieh, S.N., T. W. Rowell, D. L. Peer, and P. J. Cranford. 1991. The effects of trawling, dredging and ocean dumping on the eastern Canadian continental shelf seabed. Continental Shelf Research 11 (8—10): 1237-1263. Musick, J. A. 1999. Criteria to define extinction risk in marine fishes. Fisheries 24(12): 6-14. Moller, V., and H. J. Ratz. 1999. Assessment of Atlantic Wolffish (Anarhichas lupus L.) off west and east Greenland, 1982-98. NAFO SCR Document 99/37, series Number N4095. 14 pages. NAFO (North Atlantic Fisheries Organisation). 2000. Scientific Council Reports 2000. NAFO, Dartmouth, Nova Scotia. 164 pages. Nelson, G. A., and M. R. Ross. 1992. Distribution, growth and food habits of the Atlantic Wolffish (Anarhichas lupus) from the Gulf of Maine-Georges Bank region. Journal of Northwest Atlantic Fishery Science 13: 53-61. Pavlov, D. A., and G. G. Novikov. 1993. Life history and peculiarities of common wolffish (Anarhichas lupus) in the White Sea. ICES Journal of Marine Science 50 (3): 271-277. Powles, H. 2000. Letter and attached detailed comments from Howard Powles, Director Fisheries Research Branch, commenting on the Atlantic Wolffish draft report, January 31, 2000. Appendix A, 20 pages in O’Dea, N.R. 2001. Risk status assessment of wolffishes (Anarhichadidae) in Atlantic Canadian waters. B.Sc. (Honours) thesis, Department of Biology, Memorial University of Newfoundland, St. John’s. 99 + 20 pages. Ratz, H. J. 1997. Structures and changes of the demersal fish assemblage off Greenland, 1982-96. NAFO Scientific Council Studies 32: I-15. Schneider, D. W., T. Bult, R.S. Gregory, D. A. Methven, D. W. Ings, and V. Gotceitas. 1999. Mortality, movement, and body size: critical scales for Atlantic cod (Gadus morhua) in the northwest Atlantic. Canadian Journal of Fisheries and Aquatic Sciences 56(Supplement |): 180-187. 432 Scott, W. B., and M. G. Scott. 1988. Atlantic Fishes of Canada. Canadian Bulletin of Fisheries and Aquatic Sciences 219. 731 pages. Sinclair, A. F., and S. A. Murawski. 1997. Why have groundfish stocks declined? Pages 71-93 in Northwest Atlantic Groundfish: Perspectives on a Fishery in Collapse. Edited by J. Boreman, B. S. Nakashima, J. A. Wilson, and R. L. Kendall. American Fisheries Society. Bethesda, Maryland. Stokesbury, K. D. E., and J. H. Himmelmann. 1995. Biological and physical variables associated with aggre- gations of the giant scallop Placopecten magellanicus. Canadian Journal of Fisheries and Aquatic Sciences 52 (4): 743-753. Templeman, W. 1984. Migrations of wolffishes, Anarhichas sp., from tagging in the Newfoundland area. Journal of Northwest Atlantic Fisheries Science 5: 93-97. Templeman, W. 1986. Some biological aspects of Atlantic Wolffish (Anarhichas lupus) in the northwest Atlantic. Journal of Northwest Atlantic Fisheries Science 7 (1): 57-66. Villagarcia, M. G. 1995. Structure and Distribution of Demersal Fish Assemblages on the Northeast New- foundland/Labrador Shelf. M.Sc. thesis, Department of THE CANADIAN FIELD-NATURALIST Vol. 116 Biology, Memorial University of Newfoundland, St. John’s. 89 pages. Villagarcia, M. G., R. L. Haedrich, and J. Fischer. 1999. Groundfish assemblages of eastern Canada examined over two decades. Pages 239-259 in Fishing Places, Fishing People. Edited by D. Newell and R. E. Ommer. University of Toronto Press, Toronto. Watling, L., and E. A. Norse. 1998. Disturbance of the seabed by mobile fishing gear: A comparison to forest clearcutting. Conservation Biology 12 (6): 1180-1197. Wheeler, A. 1969. The Fishes of the British Isles and North West Europe. Michigan State University Press, East Lansing, Michigan. 613 pages. Whitehead, P. J. P., M. L. Bauchot, J.-C. Hureau, J. Nielsen, and E. Tortonese. Editors. 1986. Fishes of the North-eastern Atlantic and the Mediterranean, Unesco, Paris, volume 3. 1115 pages. Witman, J. D., and K. P. Sebens. 1992. Regional varia- tions in fish predation intensity: A historical perspective in the Gulf of Maine. Oecologia (Heidelberg) 90 (3): 305-315. Received 5 July 2001 Accepted 17 May 2002 The Muskellunge, Esox masquinongy, as a host for the Silver Lamprey, /chthyomyzon unicuspis, in the Ottawa River, Ontario/Québec CLAUDE B. RENAUD Research Division, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada Renaud, Claude B. 2002. The Muskellunge, Esox masquinongy, as a host for the Silver Lamprey, Jchthyomyzon unicuspis, in the Ottawa River, Ontario/Québec. Canadian Field-Naturalist 116(3): 433-440. A new host, Esox masquinongy, the Muskellunge, for the Silver Lamprey, Ichthyomyzon unicuspis, is reported. Fifteen Silver Lamprey/Muskellunge interactions were documented in a 90-km section of the lower Ottawa River, on the Ontario as well as the Québec side, from Ottawa to Hawkesbury between 1992 and 2001. Sites of attachment were predominantly on the back. Number of marks per host varied between 1 and 31 with a mean of 10.6. There was evidence of cytolytic activity of the buccal gland secretions. Shallowness of the fresh wounds indicated blood feeding rather than flesh feeding. Survival of the host was indicated by the presence of healed wounds in 26.7% of the cases. Muskellunge over 122 cm in total length were preferred over smaller individuals. Lampreys appeared to be more highly concentrated in the 50-km stretch of the lower Ottawa River, between Thurso and Hawkesbury, than they were in the 40-km stretch upstream, between Ottawa and Thurso. Cet article fait mention d’un nouvel héte pour la Lamproie Argentée, Jchthyomyzon unicuspis. Il s’agit du Masquinongé, Esox masquinongy. Quinze interactions entre la Lamproie Argentée et le Masquinongé furent rapportées dans une section de 90 km de la riviére des Outaouais, du cdté de |’Ontario aussi bien que celui du Québec, entre Ottawa et Hawkesbury pour la période 1992 4 2001. Les points de fixation étaient surtout sur le dos. Le nombre de marques variait entre | et 31 par héte avec moyenne 10,6. Une action cytolytique des sécrétions des glandes buccales était indiquée. Les plaies non cica- trisées étant peu profondes, une diéte de sang plut6t que de chair était indiquée. La survie de l’héte était indiquée par la présence de plaies cicatrisées dans 26,7% des cas. Les Masquinongés de taille supérieure 4 122 cm de longueur totale étaient choisit de préférence aux individus de taille moindre. Les lamproies semblaient étre plus concentrées dans une sec- tion de 50 km de la riviére des Outaouais entre Thurso et Hawkesbury qu’elles ne |’étaient dans la section de 40 km en amont, entre Ottawa et Thurso. Key Words: Silver Lamprey, Jchthyomyzon unicuspis, Petromyzontidae, parasite, Muskellunge, Esox masquinongy, Esocidae, host, Ottawa River, Ontario, Québec. Adult Silver Lampreys, [chthyomyzon unicuspis, feed ectoparasitically on the blood of a wide variety of freshwater fish species. A survey of the literature indicated that 19 freshwater fish species serve as hosts to the Silver Lamprey in nature, for feeding and/or transport (Table 1). An additional five fish species that have served as hosts under laboratory conditions (Roy 1973) are omitted rom Table 1. A record of Rainbow Trout, Oncorhynchus mykiss, acting as a host was dismissed because it was based on unconfirmed evidence, the lamprey not having been kept for identification (Hubbs and Trautman 1937). Finally, a number of other fish hosts were reported by Forbes and Richardson (1920), Starrett et al. (1960), and Hubley (1961) for Silver Lamprey and Chestnut Lamprey, and by Cochran and Marks (1995), for Silver Lamprey and Sea Lamprey; how- ever, it could not be established with certainty which species in the pair was responsible for the marks, and therefore, these were also omitted from Table 1. This paper presents data on a new host species, the Muskellunge, Esox masquinongy, based on 15 host/parasite interactions with Silver Lamprey. The Ottawa Valley Chapter of Muskies Canada has kept 43 archives, including photographs, of lamprey-induced marks on Muskellunge from the Ottawa River since 1992 and provided these to the author. The Silver Lamprey was assumed to be the only species respon- sible for the marks because it was the only one iden- tified in the seven cases in which attached lampreys were retrieved. Additionally, the only other parasitic lamprey recorded from the Ottawa River basin, the Chestnut Lamprey, Jchthyomyzon castaneus, is rare in Ontario and Québec compared to the Silver Lamprey (Renaud et al. 1996; Renaud and de Ville 2000), and therefore, is unlikely to have contributed significantly to the marks on Muskellunge reported here. Material and Methods The 15 Muskellunge in this study (Table 2) were caught by angling in the Ottawa River (Figure 1) by members of the Ottawa Valley Chapter of Muskies Canada. They were released alive after measurement of total length (TL, nearest cm), fork length (FL), and number, position (i.e., dorsal, lateral, ventral) and condition (i.e., fresh, bleeding wound or scarred, healed wound) of lamprey-induced mark(s). A 3 434 THE CANADIAN FIELD-NATURALIST a 8 Soy ea eee ee. ae eer Vol. 116 Table 1. Natural fish hosts of the Silver Lamprey, Ichthyomyzon unicuspis. Scientific and common names follow Robins et al. (1991). Fish Host Acipenser fulvescens Acipenser oxyrhynchus Polyodon spathula Lepisosteus osseus Carassius auratus Cyprinus carpio Catostomus catostomus Catostomus commersoni Ictiobus niger Ameiurus nebulosus Esox lucius Esox masquinongy Coregonus clupeaformis Salvelinus fontinalis Salvelinus namaycush Morone chrysops Morone saxatilis Ambloplites rupestris Micropterus dolomieu Stizostedion vitreum Common Name Lake Sturgeon Atlantic Sturgeon Paddlefish Longnose Gar Goldfish Common Carp Longnose Sucker White Sucker Black Buffalo Brown Bullhead Northern Pike Muskellunge Lake Whitefish Brook Trout Lake Trout White Bass Striped Bass Rock Bass Smallmouth Bass Walleye photograph of the host was taken and any attached lamprey removed and retained. In one instance, how- ever, the Muskellunge did not survive, despite attempts to release it alive, and it was retained along with the attached lamprey. The seven lamprey speci- mens and the one Muskellunge were given Canadian Museum of Nature Fish Collection (CMNFI) catalog numbers (Table 2). Total length (TL) and disc length (d) of the freshly-preserved lampreys were measured to the nearest 0.5 mm. Results The seven lampreys retrieved from the Muskel- lunge were identified as Ichthyomyzon unicuspis based on their possession of a single dorsal fin and only unicuspid endolaterals. They measured 105.0—232.0 mm TL (mean = 150.6 and standard deviation (SD) 46.6). Their oral disc lengths mea- sured 10.5—22.0 mm (mean = 14.9 and SD 4.1). The single Muskellunge kept (record number 12, Table 2) was identified using Scott and Crossman (1973). It was the only host available to the author as a spec- imen, instead of a photograph, and was therefore thoroughly examined for lamprey-induced marks. It had five fresh wounds on the dorsal surface; two immediately behind the head, one at about a third of the way down the body and two near the origin of the dorsal fin. It had one fresh wound on the right opercle and nine on the right lateral surface of the body (five fresh and four healed). It had one fresh and three healed wounds on the left opercle; the Source Vladykov (1949, 1985), Roy (1973), Becker (1983), Choudhury and Dick (1993) Vladykov (1949), Roy (1973) Wagner (1904, 1909), Hubbs and Trautman (1937), Becker (1983), Lyons (1993), Runstrom et al. (2001) Bensley (1915), this study George (1985) Becker (1983) Allin (1951), Roy (1973) Hubbs and Trautman (1937), Allin (1951), Roy (1973) Hubbs and Trautman (1937) Hubbs and Trautman (1937), George (1985) Bensley (1915), Hubbs and Trautman (1937), Becker (1983) this study Hubbs and Trautman (1937) Roy (1973) Hubbs and Trautman (1937) Becker (1983) Roy (1973) Hubbs and Trautman (1937) Hubley (1961) Hubbs and Trautman (1937) fresh one was subcircular and measured 21 mm along its longest axis. One fresh wound was present on the left branchiostegal membrane and two fresh wounds were on the left lateral surface of the body near the head. On the ventral surface it had four healed wounds: three under the right pectoral fin and one between the right pelvic and anal fin. Addi- tionally, at capture, one feeding adult lamprey was inside the branchial cavity of the host. Fourteen Muskellunge hosts measured 94—135 cm TL (mean = 122.6 and SD 12.0; the 104 cm FL Muskellunge was not included because its measure- ment was not comparable with the others). Three of the hosts (record numbers 8, 11, and 12, Table 2) weighed 18, 19, and 13 kg, respectively. The number of marks per host (n = 15) varied between 1-31 (mean = 10.6 and SD 10.5). Twelve of the 15 Muskellunge (80%) had multiple marks and two of these hosts had two attached lampreys (record num- bers 4 and 11, Table 2) at the time of capture (Figure 2). The mean number of attached lampreys on 10 hosts was 1.2. The condition of the marks (i.e., fresh or scarred) was recorded for all 15 hosts. All had fresh wounds and four of them (26.7%) also had scarred wounds. The lamprey-induced wounds, whether fresh or scarred, were more or less circular in shape and corresponded to the oral disc size. Fresh . wounds were shallow, of reddish color, and the integument over the entire surface, at the site of fresh wounds, was missing (Figure 2). The position of the lamprey-induced marks was recorded in 13 of 15 435 MUSKELLUNGE AS HOST FOR SILVER LAMPREY . . RENAUD 2002 M.97TSL N.IEoSP VN VN VN sOBJINS [BSIOP UO Ysa} ¢ (GG VN “TOATY AAQVT JO YINOU “Saqgn¢) 100z Aine 9 | soovyins OQOOT [e1oye] puv [esiop Uo MOE PSotl NSS. 8EoSP JOY Jaquiaydag VN VN VN poyeoy pue Yysolj CT -7] a ST asUOUTY JO Isk9 UI (IE INOGe ‘daqgnd) t7 vl pepsooel M.IPobl N.LEoSt VN VN VN jou uoNIsod “Yysaay 8-9 Lvl ¢ ‘[PUSLIQ,,] Jvau ‘saqgnQ/ouejUuO 0ooz Aine SI | SOOVJANS [R.IVUDA vI-1007 pue [e1oyey “[essop uo M.Ltobl N.LEoSt IHNINO O'SI O'StI poyeoy [[ pue ysody Cc] le] Vv ‘AinqsoyMeH{ vou ‘saqenQ/OuejuC 0007 Aine SI ra soBfMS [eSIOp M.CPoSL N.LCoSP €1-100C UO pe[Boy pue Ysory / ‘aSpliq Joni )-preuogor|A JO Isva 0007 IHNINOD SOI 0'SOI —9 pur [BUSA UO Ysoly | cel VN W (OZ “JOATY NROUNLD Jo YyINoUL ‘Saqand) ounf [7 I] c1-1007 pepsooel M.97SL N. 1 E0$P 0007 IHNIWO OTI O'6LI jou uontsod “ysouy | v6 VN “TOATY SAQUT JO YINOW “SaqgnC) ounf 17 Ol M.IPoSL N.LCoSt L-100Z DOVJANS [BIOL] (y)suaT] ‘BMENQC ‘AVG JOUIOAOD JO }svd “NOYOO| 6661 IHNWO gS CL8I UO peBoy 7 pur Ysa} | 104) POI 9 JO JUOAZ UL “YAW BJJIPJOOY “OLeIUO 4990190 OE 6 soovy.ins 6661 9- 1007 [eroye[ UO JoMoy TIM M.T1oSL N.SEoSh “JOAOPUa MA saquiaydag IHNIWO Oreré 0'CEC Jessop uo Apsoul Ysoy (CE eel I pur purpyooy Usemjaq ‘oaqgnd)/OlRjUC ra | 8 €- L007 M.OoSL N.9ESP IHNIWO 081 CSS SOBJINS [eSIOP UO YSolf | Sel VN ‘O]JOQoWOJ] JO ISAM “OaGgNC) 6661 Aine €1 i VN VN VN SOBJINS [P1oJe] UO YSorf ¢ vel € M.9SoPL N.6€0Sh ‘OLsqauUoW “o9qgnd 6661 Aine 9 8£-6661 M.Oo$L N.9E0SP 6661 IHNIWO cll ¢°LOI SORJINS [eSIOP UO YSoJ | COL VN ‘OJOQawUo JO ISAM “Oaqgnd) ounf 7¢ ¢ soups N.bbpobl N.LEoSh ‘IOAry Wsnoy Woy VN VN VN [esIOp UO Ysol} [¢ Lol ce ssoroe ‘Avg [PUSLIQ,"T JO OYs}JO “OLIRIUQ ~—_L66] 1990190 v soRpNs M.SSobl N.6€oSh “JOATA 9661 VN VN VN [es1op UO Ysoary O] QT Q"] ISUOUTY YIM oUINIJUOD Je ‘Sagan? snsny € X M.OE.EL0SL N..O.OoSP £661 VN VN VN ddvJ.INS [eSIOP UO YSoJJ 9 ani ¢ ‘IOATY Oyouryg 99g JO 1s9M snl ‘saqand) 1ag0199 Z SOOBJANS [PIOJC] MOE OLoSL NOE PEoSh JOATY UOHRN C661 VN VN VN pue [esiop uo Ysol} [| CTI cv YINOg JvoU “pue]Ss| [BABYS Op JJO “OLIIUO 19G019O 87 I Joquiny wu ‘p wut “TL, uOnIsOg pue uoNIpuoD wo‘T_ w ‘yideq SOJVUIPIOOD eorydessoIyH YM AVI[eI0'7T uonogJoD = Joquinyy sojeywag = =—s- Aad ure] Aoiduey = ‘syreyy Aosduiey] jo soquinyy = od unT[oyYsnyy Joye jo arg plodoy “IOATY BMLNO OY) UT SUOTORIO\UT AoIdwIe’'T JDATIS — OBuNT[aysNJY CS] IY) JOJ BIVp UONIEJOD ‘7 AIGVL * ee 436 12 Kilometres THE CANADIAN FIELD-NATURALIST Vol. 116 f pot f j MONTEBELLO 6(5)al4(12 =e SN, \ \ ie S VSS “Se a _ ONTARIO \ FIGURE 1. Co-occurrences of Muskellunge and Silver Lamprey in the Ottawa River, Ontario/Québec, 1992-2001 (solid cir- cles). The numbers without parentheses refer to the records listed in Table 2 and those in parentheses to the number of lamprey-induced marks on a Muskellunge. The locations of major centres are marked by solid triangles. hosts; 84.6% of these had marks on their dorsal sur- face, 46.2% on the lateral surface and 15.4% on the ventral surface. Six hosts had marks only on the dor- sal surface; two only on the lateral surface; and none only on the ventral surface. Three hosts had marks on both the dorsal and lateral surfaces; one had marks on both the dorsal and ventral surfaces and one had marks on all three surfaces. Muskellunge with fresh wounds were collected between 21 June and 30 October. The Muskellunge were collected at water depths between | and 6 m (mean = 3.4 and SD 1.5). During 2000, along the length of the Ottawa River which comprised the study area (Figure 1), Muskellunge less than 91 cm TL bore no marks, individuals between 91 and 122 cm TL showed a 2- 5% incidence of marks and ones greater than 122 cm TL showed a 21.4% (i.e., 6 out of 28) incidence of marks (Hedrik Wachelka, personal communication). The sample sizes for the first two categories were not reported. Discussion The adult feeding lampreys in this study were well within the range of total lengths, 85 — 392 mm, reported for adults of the species (Lanteigne 1981; Cochran and Marks 1995). Few studies have exam- ined the parasite-host interactions in the Silver Lamprey. Vladykov and Roy (1948) mentioned that transformed specimens of Silver Lamprey were col- lected on 11 species of freshwater fishes in Québec, but did not list them. George (1985) reported that the majority of about 430 dead or moribund Goldfish, Carassius auratus, all approximately 25 cm TL in a 3-km section of the Hudson River, New York State, exhibited surface or penetrative, slightly hemor- rhagic, wounds varying between 9 and 50 mm in size, and that these were attributable to Silver Lamprey. George (1985) added that the 9-mm wounds corresponded well to the disc length of the Silver Lamprey collected off the Goldfish. However, the 50-mm wounds are suggested to be caused by the Sea Lamprey, Petromyzon marinus, which also occurs in the Hudson River (Smith 1985). In the largest specimens of Silver Lamprey available to them, 301-330 mm TL, which were presumed to have the largest discs, Hubbs and Trautman (1937) 2002 RENAUD: MUSKELLUNGE AS HOosT FOR SILVER LAMPREY FIGURE 2. Muskellunge (record number 4, Table 2) with two attached feeding lampreys (visible at lower right) and 31 437 fresh lamprey-induced marks (about 20 visible) on its dorsal surface. reported a mean disc length of 7.2% of TL. If we take the maximum value in that size class range, 330 mm, this translates into a disc length of 24 mm which is slightly larger than the largest disc mea- sured in this study, 22.0 mm, but considerably less than 50 mm. Vladykov and Kott (1980) recorded a disc length of 52 mm on a 680 mm TL Sea Lamprey. Those 50-mm wounds were therefore likely caused by Sea Lamprey. The Muskellunge is an ideal host for the Silver Lamprey because of its rather small cycloid scales and the fact that as an ambush predator it lies motionless in heavily vegetated slow-moving waters waiting for prey to come within striking range. The low incidence of marks on the ventral surface of Muskellunge (15.4%) relative to the percentages which occurred on the lateral (46.2%) and dorsal (84.6%) surfaces supported the idea that while the Muskellunge lies motionless waiting for its prey, its underside was generally less available for attachment by the lamprey. The relative surface area of the three body regions of the host would not explain the dif- ference in percentage incidence of marks observed because the lateral region is at least twice (two sides) as large as either the dorsal or ventral region. This was in marked contrast with the situation reported by Wagner (1909) in Paddlefish, Polyodon spathula, where, although marks may be found on all surfaces of the fish, the attachment of Silver Lamprey was predominantly on the ventral surface. This difference in preferred site of attachment is believed to be attributable to differences in the life habits of the hosts. Unlike Muskellunge, Paddefish are pelagic fishes that are constantly moving in the water col- umn feeding on plankton (Wagner 1909), and there- fore, their underside is constantly available for attachment by lampreys. This indicates that Silver Lamprey are flexible in their site of attachment. Cochran (1986) suggested that dorsal attachments of the Chestnut Lamprey, /chthyomyzon castaneus, to hosts in shallow watercourses with relatively rough substrates were an adaptation to prevent the lam- prey’s detachment or injury through abrasion against the substrate. Similarly, protection would have been afforded the Silver Lamprey found inside the branchial cavity of one of the hosts in this study (record number 12, Table 2). Wagner (1904) report- ed at least one case in which a lamprey had found its way under the opercular flaps of Paddlefish. Because the integument was missing over the entire surface of the fresh wounds on the Muskel- lunge (Figure 2) rather than being limited to the cen- tral area of the wound where the rasping action of the transverse lingual lamina occurs, I suggest that the buccal gland secretions in /chthyomyzon unicus- pis have a cytolytic action similar to that in | Pe THE CANADIAN FIELD-NATURALIST 438 Petromyzon marinus (Lennon 1954). The shallow- ness of wounds on the Muskellunge was an indirect indication that the Silver Lamprey was a blood feed- er rather than a flesh feeder. Blood feeding in this species had been inferred by Potter and Hilliard (1987), based on its dentitionalcharacteristics, as well as its relatively large oral disc and buccal glands, being similar to that of a known blood-feeder, Petromyzon marinus. Additionally, the presence of healed injuries in 26.7% of the hosts in this study was indicative of non-lethal past events of para- sitism. Lyons (1993) stated that most of the 240 Paddlefish he examined had healed wounds and that this indicated survival from lamprey attacks. No deep wounds were found on the Muskellunge in this study. The penetrative wounds on Goldfish reported by George (1985) may well have been caused by Sea Lamprey rather than Silver Lamprey (see above). Penetrative Sea Lamprey wounds reach- ing the coelomic cavity as well as perforating cranial bones were observed in both the lab and thefield by Lennon (1954). In the lab situation, Silver Lamprey are certainly capable of penetrative wounds into the coelomic cavity and perforating cranial bones (Roy 1973), but whether this happens in nature is not known. Such severe wounding would almost certain- ly result in the relatively quick death of the host and is unlikely to be observed in nature unless large- scale sampling is conducted. The presence of fresh wounds on Muskellunge caught between 21 June and 30 October indicated that Silver Lamprey were actively feeding during that period. Authors usually do not specify whether lamprey attachments on hosts were for the purposes of feeding or transport. In the case reported here, it was quite clear from the wounds inflicted on the Muskellunge that feeding was involved. Likewise, feeding occurred in the case of Polyodon spathula (Lyons 1993) and Carassius auratus (George 1985). Although the maximum number of 28 fresh wounds in P. spathula reported by Lyons (1993) was similar to the 31 reported here for Muskellunge, the mean number of 6.1 fresh marks (Lyons 1993) was sub- stantially lower than the 9.8 (n = 13) reported here. The incidence of lamprey-induced marks in this study, 21.4% in Muskellunge larger than 122 cm TL, was significantly lower than that reported by Wagner (1909) for Paddlefish. Virtually all of the 1500 he examined from Lake Pepin, Wisconsin, had marks. Recent surveys of Paddlefish in the Wisconsin River by Lyons (1993) and Runstrom et al. (2001) have likewise reported very high incidence of Silver Lamprey marks, 96% (n = 240 Paddlefish) and 83.6% (n = 315), respectively. Although the Chest- nut Lamprey also occurs in Lake Pepin (Wagner 1909; Hubbs and Trautman 1937), it appears to be rare compared to Silver Lamprey, a situation similar to that found in the Ottawa River basin (Renaud and Vol. 116 de Ville 2000). The incidence of lamprey-induced marks in this study was positively correlated with size of the Muskellunge. Intuitively, one would think that large hosts are favored because of their large blood supply and the likelihood of recovery from an attack, making them available to future events of parasitism. It is not advantageous for a parasite to kill its host. However, in his laboratory experiments, Roy (1973) determined that Silver Lamprey attach- ments resulted in 51.9% of the hosts being killed. Additionally, he observed that the size of the host did not confer an advantage to its survival from lam- prey attacks but rather the site of attachment was the critical factor, with wounds inflicted on the head and abdomen more likely to be fatal (Roy 1973). The range in length and weight of the hosts in Roy’s (1973) experiments, however, was significantly smaller than those in this study. All species com- bined, the lengths of the hosts varied between 8-98.9 cm and weight between 0.005 and 2.2 kg, whereas the Muskellunge in this study varied between 94-135 cm TL (n = 14) and weight between 13 and 19 kg (n = 3). There might be a threshold length or weight which confers an advantage to the survival of the host that was not reached in Roy’s (1973) experiments. The cause of death of the Muskellunge in this study was not established, but a number of interesting observations were noted. It possessed 26 lamprey-induced marks, which although high were not lethal in two other cases with still higher marks. Although it possessed four marks on its ventral surface, these were healed. However, its weight was low, 13 kg for a total length of 131 cm. Furthermore, the gills on the left side were pale brown whereas the ones on the right side were dark red. One of the fishermen who was in the fishing party, Adam McLellan, reported a lamprey falling out of the branchial cavity upon landing. Perhaps, the low condition factor and the loss of blood made the Muskellunge too weak to survive the trauma of being hooked and taken out of the water. Lennon (1954) observed extensively torn gill filaments on one or more gill arches in hosts of the Sea Lamprey and suggested that the copious secretions of mucus in the branchial cavity as a result of this irritation could have impaired gill function and thereby contributed to the death of the host. No apparent damage, however, was done to the gill filaments of the Muskellunge in this study. Although Silver Lamprey has also been known to parasitize fish as small as 25-cm TL (Goldfish: George 1985), a phenomenon reported notably by Cochran and Jenkins (1994) for other lamprey species, its preference would seem to be towards _ large fish hosts (i.e., greater than 30 cm TL) as indi- cated by its size-based differential selection of Muskellunge in this study and the fact that the majority of documented hosts (14 out of 20) are 2002 RENAUD: MUSKELLUNGE AS HOST FOR SILVER LAMPREY 439 large (Table 1), based on average lengths in Scott and Crossman (1973). Very few studies actually recorded the size of the hosts. The few that did were as follows: 130 cm TL, 16 kg Acipenser fulvescens (Vladykov 1985); 97-168 cm TL Polyodon spathula (Lyons 1993); about 25 cm TL Carassius auratus (George 1985); about 84 cm TL Esox lucius (Hubbs and Trautman 1937); 94-135 cm TL E. masquinongy (this study). The five hosts considered small would be the Goldfish, the Brown Bullhead, the Brook Trout, the Rock Bass and the Smallmouth Bass. The greater incidence of marks on larger Muskellunge is suggested to be attributable to two factors: their larg- er surface area, providing more attachment sites, and their greater age, increasing the probability of encounter. In laboratory experiments, Cochran (1985) demonstrated that a congeneric lamprey, Ichthyomyzon castaneus, showed statistically signifi- cant preference for host fish with larger surface areas. Additionally, slightly over half the hosts (11 out of 20) are either naked, have large scutes con- fined to certain areas only, or in most cases possess thin scales (Table 1). The nine hosts with thick scales would be the Longnose Gar, the Goldfish, the Common Carp, the Black Buffalo, the White Bass, the Striped Bass, the Rock Bass, the Smallmouth Bass and the Walleye. The Longnose Gar as a host for the Silver Lamprey (Bensley 1915) was con- firmed in the Ottawa River (CMNFI 1999-39). Nevertheless, the typical host of the Silver Lamprey is larger than 30 cm TL and is thin scaled. Multiple marks were found on 80% of Muskellunge in this study, a level higher than that reported for Paddlefish where 70% of 315 specimens had multiple marks (Runstrom et al. 2001). Although a single lamprey may be responsible for a number of marks on a host, there were two cases in this study where two lampreys were attached to a single Muskellunge and two cases are known, one in 1999, the other in 2001, where three lampreys were attached to a Muskellunge in the 90-km section of the Ottawa River under consideration (John Anderson, personal communication). The mean number of 1.2 lamprey attached to Muskellunge (this study) was identical to that found on Paddlefish (Lyons 1993). Some Paddlefish have been found with between 10 and 25 Silver Lampreys attached (Wagner 1909; Lyons 1993). The report by Vladykov (1985) of 61 Silver Lamprey on a Lake Sturgeon is an extreme case of multiple lampreys on a host. It is known that Muskellunge outside the spawning season (late April — early May) are both solitary and sedentary (Scott and Crossman 1973). Our material was collected during June-October, thus post-spawning. Therefore, by mapping the loca- tion of the Muskellunge with high numbers of lam- prey marks, arbitrarily set at 10 marks, it appears that adult Silver Lamprey are more highly concen- trated in a 50-km stretch along the lower Ottawa River, between Thurso and Hawkesbury, compared to the 40-km stretch upstream, between Ottawa and Thurso (Figure 1). Indeed, in the former case there are ten records, six of which have over ten marks (overall mean = 13.9), while in the latter there are only five records, all below ten marks (mean = 4.0). Acknowledgments The author expresses his thanks to the following members of the Ottawa Valley Chapter of Muskies Canada: John Anderson, Greg Bright, Rick Collins, Marc Dunn, Sean Hanrahan, Ed Lalonde, Charlie Landreville, Jeff Little, Adam and Stefan McLellan, Ken Taggart, Denise and Hedrik Wachelka (the President). Noel Alfonso, Canadian Museum of Nature, produced figure 1 and Sylvie Laframboise, of the same institution, helped in the lab. Two anonymous reviewers provided much appreciated and helpful suggestions. Literature Cited Allin, A. E. 1951. Records of the Sea Lamprey and the Silver Lamprey from Canadian waters of the western end of Lake Superior. Canadian Field-Naturalist 65: 184-185. Becker, G. C. 1983. Fishes of Wisconsin. The University of Wisconsin Press, Madison, Wisconsin. xii + 1052 pages. Bensley, B. A. 1915. The fishes of Georgian Bay. Contributions to Canadian Biology Sessional Paper 39b, Fasc. II: 1-51 + 6 figures + 2 plates. Choudhury, A., and T. A. Dick. 1993. Parasites of Lake Sturgeon, Acipenser fulvescens (Chondrostei: Acipenser- idae), from central Canada. Journal of Fish Biology 42: 571-584. Cochran, P. A. 1985. Size-selective attack by parasitic lampreys: consideration of alternate null hypotheses. Oecologia 67: 137-141. Cochran, P. A. 1986. Attachment sites of parasitic lam- preys: comparisons among species. Environmental Biology of Fishes 17: 71-79. Cochran, P. A., and R. E. Jenkins. 1994. Small fishes as hosts for parasitic lampreys. Copeia 1994: 499-504. Cochran, P. A., and J. E. Marks. 1995. Biology of the Silver Lamprey, Ichthyomyzon unicuspis, in Green Bay and the Lower Fox River, with a comparison to the Sea Lamprey, Petromyzon marinus. Copeia 1995: 409-421. Forbes, S. A., and R. E. Richardson. 1920. The fishes of Illinois. Natural History Survey of Illinois 3: cxxxvi + 357 pages. George, C. J. 1985. Occurrence of the Silver Lamprey in the Stillwater sector of the Hudson River. New York Fish and Game Journal 32: 95. Hubbs, C. L., and M. B. Trautman. 1937. A revision of the lamprey genus /chthyomyzon. Miscellaneous Publications of the Museum of Zoology, University of Michigan (35): 7—109 + 2 plates. Hubley, R. C., Jr. 1961. Incidence of lamprey scarring on fish in the Upper Mississippi River, 1956-58. Transactions of the American Fisheries Society 90: 83-85. a Se 440 Lanteigne, J. 1981. The taxonomy and distribution of the North American lamprey genus Jchthyomyzon. M.Sc. thesis, University of Ottawa, Ottawa, Canada. xi + 150 pages. Lennon, R. E. 1954. Feeding mechanism of the Sea Lamprey and its effect on host fishes. Fishery Bulletin 56: 247-293. Lyons, J. 1993. Status and biology of Paddlefish (Polyodon spathula) in the lower Wisconsin River. Transactions of the Wisconsin Academy of Sciences, Arts and Letters 81: 123-135. Potter, I. C., and R. W. Hilliard. 1987. A proposal for the functional and phylogenetic significance of differ- ences in the dentition of lampreys (Agnatha: Petromyzontiformes). Journal of Zoology, London 212, Part 4: 713-737. Renaud, C. B., and N. de Ville. 2000. Three records of the Chestnut Lamprey, /chthyomyzon castaneus, new to Québec. Canadian Field-Naturalist 114: 333-335. Renaud, C. B., S. C. Ribey, and F. Chapleau. 1996. Four records of the Chestnut Lamprey, /chthyomyzon castaneus, new to Ontario. Canadian Field-Naturalist 110: 450-453. Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott. 1991. Common and scientific names of fishes from the United States and Canada. American Fisheries Society Special Publication 20: 1-183. Roy, J.-M. 1973. Croissance, comportement et alimenta- tion de la lamproie du nord (Ichthyomyzon unicuspis, Hubbs et Trautman) en captivité. Travaux sur les Pécheries du Québec (41): 3-144. Runstrom, A. L., B. Vondracek, and C. A. Jennings. 2001. Population statistics for Paddlefish in the Wiscon- Mm Sf och eee me et, pg hee THE CANADIAN FIELD-NATURALIST Vol. 116 sin River. Transactions of the American Fisheries Society 130: 546-556. Scott, W. B., and E. J. Crossman. 1973. Freshwater fish- es of Canada. Fisheries Research Board of Canada Bul- letin 184: xi + 966 pages. Smith, C. L. 1985. The inland fishes of New York State. The New York State Department of Environmental Conservation, Albany, New York. xi + 522 pages. Starrett, W. C., W. J. Harth, and P. W. Smith. 1960. Parasitic lampreys of the genus Jchthyomyzon in the rivers of Illinois. Copeia 1960: 337-346. Vladykov, V. D. 1949. Quebec lampreys (Petromyzoni- dae). I- List of species and their economical importance. Contribution au Département des Pécheries, Québec (26): 7-67. Vladykov, V. D. 1985. Record of 61 parasitic lampreys (Ichthyomyzon unicuspis) on a single sturgeon (Aci- penser fulvescens) netted in the St. Lawrence River (Québec). Naturaliste canadien (Revue d’écologie et de systématique) 112: 435-436. Vladykov, V.D., and E. Kott. 1980. Description and key to metamorphosed specimens and ammocoetes of Petro- myzonidae found in the Great Lakes Region. Canadian Journal of Fisheries and Aquatic Sciences 37: 1616-1625. Vladykov, V. D., and J.-M. Roy. 1948. Biologie de la lam- proie d’eau douce (Ichthyomyzon unicuspis) aprés la méta- morphose. Revue Canadienne de Biologie 7: 483-485. Wagner, G. 1904. Notes on Polyodon, I. Science (n.s.) 19: 554-555. Wagner, G. 1909. Notes on the fish fauna of Lake Pepin. Transactions of the Wisconsin Academy of Sciences, Arts and Letters 16, Part 1: 23-37. Received 9 July 2001 Accepted 23 April 2002 Ephemeral Occurrence of the Mosquito Fern, Azolla caroliniana, at Ottawa, Ontario STEPHEN J. DARBYSHIRE Agriculture and Agri-Food Canada, Eastern Cereal and Oilseed Research Centre, Wm. Saunders Building #49, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada Darbyshire, Stephen J. 2002. Ephemeral occurrence of the Mosquito Fern, Azolla caroliniana, at Ottawa, Ontario. Canadian Field-Naturalist 116(3): 441-445. Mosquito fern, Azolla caroliniana, is known in Canada only from records at Hamilton (1862) and Gananoque (1981), where it did not persist. In 1997 it was found in a pond in a residential area of central Ottawa. In 1998 is was not detected at this pond, but was found in the Rideau River through central Ottawa and into the Ottawa River on both the Ontario and Quebec shores. The enormous populations which had spread in the rivers in 1998 were completely absent in 1999. Key Words: Azolla caroliniana, Mosquito Fern, invasive plants, weeds, rare plants, plant dispersal. The fern family Azollaceae contains one genus, Azolla, with about five to seven extant species and a number of others known from fossil material. Commonly called mosquito ferns, they are small free-floating aquatic plants distributed in tropical and warm temperate regions throughout the world. Morphological reduction is exhibited along with various adaptations to its aquatic habitat. Copiously branched short stems bear alternate and (usually) closely imbricate leaves and a few pendulous roots. The leaves are divided into two lobes, a slightly larger, thin and translucent submersed lobe and a green or reddish emersed lobe covered with minute papillae. Three species are native to North America with two reported in Canada, Azolla mexicana C. Pres| in Brit- ish Columbia, and A. caroliniana Willd. in Ontario (Cody and Britton 1989). Another species, A. filicu- loides Lam. has been reported for southern Alaska (Svenson 1944; Lumpkin 1993) and may possibly be found in British Columbia. The distribution maps for A. mexicana and A. caroliniana in Cody and Britton (1989) have the captions inadvertently switched. Widespread in the eastern United States, south through Central America and the West Indies to South America, A. caroliniana has also been intro- duced to Europe and Asia (Lumpkin 1993). In spite of its wide distribution, Lellinger (1985) considered it a “rare” plant. Other authors report it as common or even abundant, but, because of its rapid growth under suitable conditions, large populations can cover connected water bodies quickly, giving an illu- sion of widespread abundance. Although known from northern New York State (see overview in Cody and Schueler 1988), it has been found only twice in Ontario (Cody and Schueler 1988). The first collection in Canada is of plants taken at Burlington Beach (near Hamilton) in 1862. The plant does not seem to have persisted there, and has not been seen since the original collection. An extensive popula- tion found at Knights Creek (near Gananoque) in 1981 also did not persist to the following year (Cody and Schueler 1988). Azolla caroliniana is a plant of slow-moving or stagnant water in ponds, lakes, marshes, swamps, streams, rivers, ditches, etc. (Lumpkin 1993). It is the most cold resistant species in the genus (Lumpkin 1993), and Redman (1995) reported that populations successfully over-wintered in Maryland and Wash- ington, D.C. Studies on physiology and responses to environmental conditions have mostly been conducted on the economically important, tropical and subtropi- cal species A. pinnata R. Br. and are summarized by Wagner (1997). Species of Azolla are heterosporic, but reproduc- tion in A. caroliniana seems to be entirely vegetative. Megaspores are unknown, although microspores are sometimes detected and are produced under condi- tions of crowding (Lellinger 1985). A symbiotic relationship has formed between species of Azolla and the cyanobacterium (blue- green alga) Anabaena azollae Strasb. (and possibly other bacteria) much like the relationship between the Fabaceae (Leguminosae) and Rhizobium (Wagner 1997). The cyanobacterium is able to fix atmospheric nitrogen, thus providing a source of this nutrient normally unavailable to plants (Wagner 1997). Although nitrogen is not a limiting nutrient, phosphorus availability can be a major constraint on Azolla growth (Kushari and Watanabe 1991; Wagner 1997). Reports of minimal phosphorous concentra- tions required for maximum growth vary between studies as well as between species. Concentrations ranging from 400 pg |! to 20,000 pg I-' have been reported as minimum requirements for maximum growth rates. Under optimal conditions some Azolla species can double their biomass in 3—5 days (Lumpkin and Plunknett 1982; Wagner 1997). 44] 442 Because of the high nitrogen content, various species are used in agriculture as a green manure or biofertilizer (Lumpkin and Plucknett 1982). Some species, including A. caroliniana, are used as orna- mental plants in aquaria and decorative pools (Lellinger 1985; L. H. Bailey Hortorium 1976). Other uses and applications are summarized by Wagner (1997). In some situations A. caroliniana has been a serious weed disrupting aquatic ecology and interfer- ing with human activities (Thieret 1980). The reduc- tion of light (up to 90%) and reduced oxygen levels (50% or more) in the water column under Azolla mats can have such an affect on other vegetation that it has been studied as a means of weed control in rice pad- dies (Wagner 1997). Azolla caroliniana at Ottawa In September of 1997.a population of A. carolini- ana was found in the westerly pond of Brown’s Inlet off the Rideau Canal in Ottawa, Ontario (Figure 1, site 1; Appendix 1, specimen 1). Individual plants and mats in patches up to about 0.5 m in diameter were common around the perimeter of the pond among other floating and emergent plants. Other vegetation studies in this pond during the previous two years had failed to detect the fern (personal observations). Kettle Island 45° 28" 45° 24’ 75° 43° 75° 41' FIGURE |. Map of the area of the Rideau and Ottawa Rivers where Azolla caroliniana populations occurred in 1997 and 1998. Numbers correspond to approxi- mate locations of collections listed in Appendix 1. THE CANADIAN FIELD-NATURALIST Vol. 116 The pond, being connected to the Rideau Canal system, is completely drained in the autumn (usually October) of each year. Draw-down of water in the Canal commenced on 15 October 1997. Drainage of the pond was complete by 18 October and mats of A. caroliniana were left stranded on the wet mud. On 19 and 20 October, daily minimum temperatures were about -3°C. The first significant snowfall occurred on October 21 (9 cm) and then again on October 26 (18 cm); however, persistent snow cover did not occur until November 15. During this time minimum temperatures of less than -5°C were recorded on six days (24, 26 and 27 October, and 12, 13 and 14 November), including the minimum of -9.4°C on 13 November. Stranded plants would have had little protection from dessication and cold tem- peratures for more than a month. Continued observa- tions during the next two years did not detect A. car- oliniana at Brown’s Inlet. It is presumed that these winter conditions proved too harsh for the vegetative material to over-winter. In late summer of 1998, colonies of A. caroliniana were seen in substantial mats along the shores of the Rideau River. Through the month of September sur- veys were conducted along the Rideau River and the Ottawa River downstream from the mouth of the Rideau River. Although a much wider area was sur- veyed (upstream on the Rideau River to Manotick, downstream on the Ottawa River to Cumberland, and the Rideau Canal from Mooney’s Bay to the Ottawa River), A. caroliniana was found only between collec- tion sites 7 and 9 (Figure 1; Appendix 1). Over this stretch of about 10 kilometres it was abundant, often completely swathing the shore for considerable dis- tances. Mats became sparser at the upstream and downstream extremes of the distribution and where the river shores were exposed to strong currents or waves. Mats were mostly along shores and becoming thick and continuous in quiet bays and around protect- ing emergent vegetation such as Cattails (Typha lati- folia L.) or Reed Canary Grass (Phalaris arundinacea L.). Sometimes mats would become dislodged in cur- rents and drift downstream. In fast running water the mats would disintegrate into individual plants or small clumps, but in slow moving water contiguous mats of about a hundred square metres were sometimes seen. While not at optimum growth levels, phosphorous concentrations measured every two weeks in the Rideau River showed a plentiful supply of this nutri- ent as well as a peak during the major population expansion in mid-July, 1998 (Figure 2). Systematic and repeated searches in August and September of 1999 throughout the area of the previ- ous year’s infestation failed to detect a single plant of A. caroliniana. Less detailed searches in 2000 were also negative. The large population seems to be the result of a single introduction at a point close to collection site number 9 (Figure 1). Figure 2 2002 DARBYSHIRE: OCCURRENCE OF MOSQUITO FERN AT OTTAWA 443 =P 1998 = # SRP 1998 —%- TP 1999 — * — SRP 1999 bi-weekly sample points (starting May 12th) FIGURE 2. Plot of the total phosphorus (TP) and soluble reactive phosphorus (SRP) in the Rideau River during the summers of 1998 and 1999 (Paul Hamilton, personal com- munication). Samples were taken at 2 week intervals starting May 12 at station #103 (St. Patrick Street Bridge), about half way between points 2 and 5 on Figure 1. indicates that phosphorous concentrations would not have been a limiting factor in 1999, so the complete disappearance is most likely the result of winter-kill. Dispersal of Azolla The west bank of the Rideau River at collection site 9 has a gentle slope allowing easy access to the water and is immediately adjacent to a small residen- tial enclave. The introduction could have easily occurred here from the dumping of aquarium con- tents. Species of Azolla are easily and often grown in aquaria for decorative purposes and used in science teaching to demonstrate the nitrogen fixing hetero- cysts of the symbiotic cyanobacteria. Small (1935) considered the occurrences of A. caroliniana around southeastern New York State and northern New Jersey (and by implication further north) to be colonies naturalized from human introductions. Fernald (1950) also considered northern occurrences as being likely spread from cultivation. In recent years a number of exotic aquarium-trade species have been reported as introduced into the Rideau Canal and Rideau River round Ottawa, including several tropical fish species (Renaud and Phelps 1999; Renaud and Phelps 2001) and a turtle (Darbyshire 2000). It may well be that the introduc- tion of A. caroliniana into the Rideau River occurred coincidental with the dumping of fish or other aquar- ium animals early in season of 1998. The occurrence of the plant in Brown’s Inlet (collection 1, Appendix 1) was most likely a deliberate introduction. Brown’s Inlet consists of two ponds in an urban resi- dential area. They are separated from each other and from the Rideau Canal by roadbeds, but all connect- ed by submerged culverts. Azolla was found only in the upper pond which has a small municipal park on the south side and residential property to the north. Other exotic plants sold for aquaria and ponds have also been collected in this same pond. Some of these have established and persisted for many years (e.g., Nymphoides peltata (Gmel.) Kuntze), while others have been ephemeral (e.g., Pistia stratiotes L.). Cranfill (1980), observing the ephemeral nature of populations and the species’ North American dis- tribution, speculated that A. caroliniana is continual- ly re-introduced by migrating waterfowl. The most upstream site of the 1998 infestation, collection site 9, is immediately downstream from a shallow area in the Rideau River which attracts large numbers of ducks. Extensive parks with ready shore access on the east side of the river (especially immediately upstream) are popular areas for people to feed water- fowl generating an additional attraction for birds. Ridley (1930) made some very interesting obser- vations on the various actual and probable distribu- tion vectors of Azol/a species in Europe and Great Britain. In addition to river flow and flood water, he cited water-birds, frogs, and human activity as —————————————————————————E——E—E———— 4 dispersal agents. Ridley speculated that birds are the most likely vehicle for long distance travel and explanation for its presence in isolated ponds. As indirect evidence he cites the sudden appearance of Azolla in a small isolated pond about 0.25 miles from a larger pond in which it was abundant. Moorhens (Gallinula chloropus) were seen going from the larger pond to the smaller one and he claimed that this would have been the only means of transport to the small pond. He also cites a fascinat- ing series of observations by Chateau where “toads” and/or “frogs” were seen to transport A. caroliniana out of pans where the fern was cultivated. Subse- quently it appeared in two nearby ponds, one of which was said to be 200 metres away. Although the fronds are reported to be able to with- stand hard frosts (-5°C) and prolonged ice cover (Lumpkin 1993), Azolla caroliniana did not persist over the winter at Ottawa. No plants were seen during the summer of 1998 at Brown’s Inlet. No plants were seen anywhere in the Rideau or Ottawa Rivers in 1999 or 2000, in spite of surveys throughout the growing season and the area of previous infestation. It is not known what mechanism brought A. caroliniana to the Ottawa area in two separate years at two sepa- rate locations. It is possible that it arrived early in the growing season transported from more southerly pop- ulations by migrating waterfowl. Circumstantial evi- dence and the prevalence of recent reports of liberated aquarium fauna and flora suggest that human agents are the most likely cause, especially in the case of Brown’s Inlet. Acknowledgements T. A. Lumpkin kindly confirmed the identification of Azolla caroliniana. Susan Flood provided the map in Figure 1. Paul Hamilton (Canadian Museum of Nature) provided data on phosphorus levels at sta- tion number 103 on the Rideau River (Figure 2). Weather data from the Ottawa CDA climate refer- ence station (CRS) jointly operated by Environment Canada and Agriculture and Agri-food Canada (45° 23'N, 75°43'W). J. Cayouette, W. J. Cody, Lynn Gillespie and L.V. Hills made useful sugges- tions on earlier versions of the manuscript. Literature Cited Cody, W. J., and D. M. Britton. 1989. Ferns and fern allies of Canada. Agriculture Canada, Research Branch Publication 1829/E. 430 pages. Cody, W. J., and F. W. Schueler. 1988. A second record THE CANADIAN FIELD-NATURALIST = | Vol. 116 - of the Mosquito Fern, Azolla caroliniana, in Ontario. Canadian Field-Naturalist 102: 545-546. Cranfill, R. 1980. Ferns and fern allies of Kentucky. Kentucky Nature Preserves Commission Scientific and Technical Series Number 1. 284 pages. Darbyshire, S. J. 2000. More red-eared sliders in the Ottawa District. Trail & Landscape 34: 15-16. Fernald, M. L. 1950. Gray’s manual of botany. A hand- book of the flowering plants and ferns of the central and northeastern United States and adjacent Canada. 8" edi- tion. American Book Company, New York, Cincinnati, Chicago, Boston, Atlanta, Dallas, San Francisco. 1632 pages. Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index Herbariorum. Part I: The herbaria of the World. 8% edition. Regnum Vegetabile 120. New York Botanical Garden, Bronx, New York. 693 pages. Kushari, D. P., and I. Watanabe. 1991. Differential responses of Azolla to phosphorus deficiency. I. Screen- ing methods in quantity controlled-condition. Soil Science and Plant Nutrition 37: 271-282. Lellinger, D. B. 1985. A field manual of the ferns & fern- allies of the United States & Canada. Smithsonian Institution Press, Washington, D.C. 389 pages. L. H. Bailey Hortorium. 1976. Hortus Third. Macmillan, New York. 1290 pages. Lumpkin, T. A. 1993. Azollaceae Wettstein. Azolla Family. Flora of North America North of Mexico. Flora of North America editorial committee. Oxford Univer- sity Press, New York, Oxford. 2: 338-342. Lumpkin, T. A., and D. L. Plucknett. 1982. Azolla as a green manure: use and management in crop production. Westview tropical agriculture series Number 5. West- view Press, Boulder, Colorado. 230 pages. Redman, D. E. 1995. Noteworthy collections. Castanea 60: 82-84. Renaud, C. B., and A. Phelps. 1999. Oscar-winning catch. Trail & Landscape 33: 178-180. Renaud, C. B., and A. Phelps. 2001. A pacu/piranha in the Rideau Canal. Trail & Landscape 35: 86-89. Ridley, H. N. 1930. The dispersal of plants through the world. L. Reeve & Co., Ashford, Kent, U. K. 744 pages. Small, J. K. 1935. Ferns of the vicinity of New York. Being descriptions of the fern-plants growing naturally within a hundred miles of Manhattan Island. With Notes. The Science Press, Lancaster, Pennsylvania. 285 pages. Svenson, H. K. 1944. The New World species of Azolla. American Fern Journal 34: 69-84. Thieret, J. W. 1980. Louisiana ferns and fern allies. Lafayette Natural History Museum, Lafayette, Louisi- ana. 124 pages. Wagner, G. M. 1997. Azolla: a review of its biology and utilization. The Botanical Review 63: 1-26. Received 24 October 2001 Accepted 22 May 2002 2002 DARBYSHIRE: OCCURRENCE OF MOSQUITO FERN AT OTTAWA 445 APPENDIX 1. Voucher specimens of Azolla caroliniana Willd. collected around Ottawa 1997-1998. A complete set is deposited at the herbarium DAO (Holmgren et al. 1990) with duplicates of some collections distributed to other herbaria. Since 1998 the Ottawa-Carleton Regional Municipality has became part of the City of Ottawa and now is included in the latter name. 1. Ontario, Ottawa-Carleton Regional Municipality, Ottawa, Brown’s Inlet, 45°23’55”N, 75°41'28”’W, artifi- cial pond along Rideau Canal, mats forming among patches with Nymphoides peltata, S.J. Darbyshire 4748, 7 September 1997. 2. Ontario, Ottawa-Carleton Regional Municipality, Ottawa, Green Island, Rideau River, 45°26’23’N, 75°41'28”W, common in shallow water among other floating/emergent vegetation, S. J. Darbyshire & P. Ouellet 4984, 30 August 1998. 3. Ontario, Ottawa-Carleton Regional Municipality, Ottawa River, Governor Bay, below 24 Sussex Drive, 45°26'45"N, 75°41'27"W, large mats in shallow bay, S. J. Darbyshire, M. Murray & Tracey McDonald 4985, 10 September 1998. 4. Ontario, Ottawa-Carleton Regional Municipality, Ottawa River across from Gatineau (Quebec), Rockcliffe Park Boathouse, 45°27'18"N, 75°41'16”W, common in sheltered waters along shore, S. J. Darbyshire, M. Murray & Tracey McDonald 4986, 10 September 1998. 5. Ontario, Ottawa-Carleton Regional Municipality, Vanier, Cummings Bridge over Rideau River, 45°25'54"N, 75°40’ 15"W, S. J. Darbyshire, M. Murray & Tracey McDonald 4987, 10 September 1998. 6. Quebec, Hull, Leamy Lake outflow and Ottawa River, 45°26'57”N, 75°42'19"W, shallow, quiet water near mouth of creek, common along river shore in quiet water, S. J. Darbyshire & M. Murray 4998, 29 September 1998. 7. Quebec, Gatineau, Ottawa River, near the west end of Kettle Island, 45°27’51”N, 75°40’55”W, shallow water among emergent vegetation, with Scirpus, Pontederia, Sagittaria, Lemna, Typha, S. J. Darbyshire & M. Murray 5000, 29 September 1998. 8. Ontario, Ottawa-Carleton Regional Municipality, Ottawa, Hurdman Bridge, Rideau River, 45°24'59’N, 75°39'53"W, large patches around Phalaris arundinacea and emergent vegetation, with Spirodella, S. J. Darbyshire & M. Murray 5001, 29 September 1998. 9. Ontario, Ottawa-Carleton Regional Municipality, Ottawa, near George Mcllraith Bridge, Rideau River, 45°24'07"N, 75°40'10"W, shallow water along shore with emergent vegetation, S. J. Darbyshire & M. Murray 5002, 29 September 1998. An additional specimen is at the Canadian Museum of Nature (CAN). This collection was made about halfway between sites 2 and 5 in Figure 1. Ontario, Ottawa-Carleton Regional Municipality, Rideau River at the St. Patrick Street bridge, east bank just south of bridge, 45°26.2'N, 75°40.5’W, band of emergent and subemergent vegetation in area with mod- erate flow, hard gravel/mud substrate; small green and red floating plants, forming a mat along shore over a large area; tiny leaves, roots hanging 5-10 cm, M Richard & R. Boles 98-217, 30 September 1998. . T_T im MMe New Records of Vascular Plants in the Yukon Territory IV WILLIAM J. Copy!, CATHERINE E. KENNEDY2, BRUCE BENNETT?, and VALERIE LOEWEN? \Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada 2Department of Renewable Resources, Government of the Yukon, Box 2703, Whitehorse, Yukon Y1A 2C6 Canada William J. Cody, Catherine E. Kennedy, Bruce Bennett, and Valerie Loewen. 2002. New records of vascular plants in the Yukon Territory IV. Canadian Field-Naturalist 116(3): 446-474. Two native taxa, Crassula aquatica and Erysimum inconspicuum, and two introduced taxa, Elymus elongatus ssp. ponticus and Erysimum cheiri, are reported new to the known flora of the Yukon Territory. Significant range extensions for 199 native and 27 introduced taxa and comments on one native taxon, Ranunculus cooleyae, previously reported in the Territory but omitted in the Flora, are also included. Key Words: Vascular plants, Yukon Territory, flora, new records, range extensions, phytogeography. Since the writing of New Records of Vascular Plants in the Yukon Territory III (Cody et al. 2001), a considerable number of plant specimens have been submitted to Cody for identification and confirma- tion, in particular from Bruce Bennett, while work- ing for Yukon Department of Renewable Resources as a wildlife biologist volunteering for the Canadian Parks and Wilderness Society on the Wind River and in various other areas in the Territory, and Valerie Loewen while studying the vegetation adjacent to the Bonnet Plume River. In addition, the senior author spent three weeks with David Cody surveying flora adjacent to the highways in the southern part of the Territory. This paper serves to further update the Flora of the Yukon Territory (Cody 1996) and Flora of the Yukon Territory, Second Edition (Cody 2000) along with other additional records recently published (Cody et al. 1998, 2000, 2001). The floristic infor- mation presented earlier and updated here is essen- tial for biological research and ongoing work relat- ing to agriculture, forestry, sustainable resource management and wildlife management. With addi- tions of two native and two introduced species reported here the flora now includes 1163 species. The new native species are rare (as defined by Douglas et al. 1981) in the Territory. The taxa addressed in the body of this paper appear first in a synoptic list by Yukon status in alphabetical order. The taxa are then discussed in taxonomic order, as presented in the Flora of the Yukon Territory with citation of specimens and other pertinent information. Common names follow Cody (1996) and Douglas et al. (1984, 1990, 1991, 1998). Two native taxa, Crassula aquatica and Erysimum inconspicuum and two introduced taxa, Elymus elon- galus ssp. ponticus, Erysimum cheiri are reported new to the flora of the Yukon Territory. Significant range extensions within the Territory are reported for 199 native and 27 introduced taxa and comments are presented on one native taxon, Ranunculus cooleyae. Synoptic List by Yukon Status Native taxa new to the Yukon Territory (2) Crassula aquatica Erysimum inconspicuum Introduced taxa new to the Yukon Territory (2) Elymus elongatus ssp. ponticus Erysimum cheiri Range extensions of native taxa within the Yukon Territory (199) Achillea millefolium ssp. lanulosa Achillea sibirica Allium schoenoprasum ssp. sibiricum Amelanchier alnifolia Amerorchis rotundifolia Androsace chamaejasme ssp. lehmanniana Antennaria monocephala ssp. monocephala Antennaria pulcherrima Arabis columbiana Arabis holboellii var. secunda Arabis nuttallii Arabis pinetorum Arctostaphylos rubra Arctostaphylos uva-ursi Arenaria longipedunculata Armeria maritima ssp. arctica Artemisia alaskana Artemisia campestris Artemisia michauxiana Artemisia tilesii Aster sibiricus Astragalus alpinus Astragalus australis Astragalus bodinii Astragalus williamsii Betula glandulosa Betula neoalaskana Boykinia richardsonii -Bromus pumpellianus var. arcticus Calamagrostis canadensis ssp. langsdorfii 446 | ee 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV Calamagrostis stricta ssp. stricta Calla palustris Calypso bulbosa Campanula aurita Campanula uniflora Cardamine bellidifolia Cardamine digitata Cardamine pratensis Carex aquatilis ssp. aquatilis Carex concinna Carex gynocrates Carex lasiocarpa ssp. americana Carex lugens Carex misandra Carex nardina Carex pellita Carex podocarpa Carex rupestris Carex saxatilis Carex scirpoidea Carex spectabilis Carex tenuiflora Carex vaginata Chamaedaphne calyculata Coeloglossum viride ssp. bracteatum Corallorhiza trifida Cornus canadensis Cornus stolonifera Crepis nana Cypripedium guttatum Cypripedium parviflorum var. makasin Cystopteris fragilis Cystopteris montana Descurainia incisa vat. incisa Draba alpina Draba borealis Draba stenoloba Elaeagnus commutata Eleocharis uniglumis Elymus calderi Elymus glaucus Elymus macrourus Elymus trachycaulus ssp. glaucus Elymus trachycaulus ssp. novae-angliae Equisetum fluviatile Equisetum palustre Equisetum pratense Equisetum scirpoides Erigeron hyssopifolius Erigeron lonchophyllus Eriophorum angustifolium Eriophorum vaginatum Erysimum cheiranthoides Festuca baffinensis Festuca brachyphylla Festuca lenensis Festuca vivipara ssp. glabra Geocaulon lividum Geum aleppicum ssp. strictum Geum macrophyllum ssp. perincisum Goodyera repens Gymnocarpium jessoense ssp. parvulum Halimolobus mollis Haplopappus macleanii Hierochloe hirta ssp. arctica Hordeum jubatum Isoetes echinospora Juncus alpinoarticulatus ssp. americanus Juncus balticus var. alaskanus Juncus castaneus ssp. castaneus Juncus castaneus ssp. leucochlamys Juncus triglumis ssp. albescens Juniperus communis ssp. depressa Juniperus horizontalis Ledum groenlandicum Lepidium bourgeauanum Lesquerella arctica ssp. calderi Linnaea borealis ssp..americana Luzula arctica ssp. arctica Luzula confusa Luzula parviflora ssp. parviflora Luzula rufescens Luzula spicata Lycopodium annotinum var. pungens Lycopodium clavatum var. monostachyon Lycopodium selago Menyanthes trifoliata Mertensia paniculata var. paniculata Minuartia biflora Minuartia rossti Moehringia lateriflora Moneses uniflora Nuphar polysepalum Orthilia secunda Oxycoccus microcarpus Oxytropis campestris ssp. varians Parrya arctica Parrya nudicaulis Pedicularis capitata Pedicularis langsdorfii ssp. arctica Pedicularis verticillata Petasites frigidus ssp. frigidus Petasites frigidus ssp. palmatus Phlox alaskensis Picea mariana Pinus contorta ssp. latifolia Plantago eriopoda Platanthera obtusata Poa abbreviata ssp. abbreviata Poa alpina Poa arctica ssp. arctica Poa glauca Polemonium boreale Polygonum buxiforme Polygonum viviparum Populus balsamifera ssp. balsamifera Populus tremuloides Potamogeton friesti Potamogeton richardsonit Potentilla anserina Potentilla litoralis Potentilla norvegica Potentilla palustris Primula mistassinica Pyrola asarifolia Pyrola chlorantha Pyrola grandiflora Ranunculus cymbalaria Ranunculus nivalis Ribes hudsonianum 447 9 Ra I 6 is 1 og 448 THE CANADIAN FIELD-NATURALIST Vol. 116 Rorippa barbareifolia Thlaspi arvense Rorippa palustris ssp. palustris Trifolium hybridum Rubus arcticus ssp. acaulis Trifolium pratense Rubus idaeus Sagittaria cuneata Salicornia europaea Salix arbusculoides Salix barrattiana Salix myrtillifolia Salix planifolia ssp. pulchra Salix pseudomonticola Salix rotundifolia ssp. dodgeana Sanguisorba officinalis Saxifraga lyallii ssp. hultenii Saxifraga nivalis Saxifraga serpyllifolia Scheuchzeria palustris ssp. americana Scutellaria galericulata var. pubescens Selaginella selaginoides Senecio atropurpureus ssp. frigidus Shepherdia canadensis Silene acaulis ssp. subacaulescens Sium suave Solidago canadensis var. salebrosa Solidago simplex Sparganium minimum Sparganium multipedunculatum Spiraea beauverdiana Spiranthes romanzoffiana Stellaria longipes Stipa hymenoides Taraxacum ceratophyllum Tofieldia coccinea Triglochin maritimum Trisetum sibiricum ssp. sibiricum Trisetum spicatum Utricularia vulgaris ssp. macrorhiza Viburnum edule Wilhelmsia physodes Range extensions of introduced taxa within the Yukon Territory (27) Alopecurus pratensis Artemisia biennis Brassica campestris Bromus inermis Capsella bursa-pastoris Chenopodium album Festuca rubra Festuca trachyphylla Lepidium ramosissimum Matricaria matricarioides Matricaria perforata Melilotus alba Melilotus officinalis Onobrynchis viciifolia Phalaris arundinacea Phleum pratense Plantago major Poa annua Poa pratensis ssp. pratensis Rheum rhaponticum Rumex crispus Sisymbrium altissimum Taraxacum officinale Trifolium repens Comments on native taxa in the Yukon Territory (1) Ranunculus cooleyae Annotated Species List LYCOPODIACEAE Lycopodium annotinum L. var. pungens (La Pylaié) Desv., Bristly Club-moss — YUKON: open disturbed ATV trail, Wind River, McClusky Lake, 64°34.19’N 134°25.77'W, B. Bennett 00-244, 2 July 2000 (DAO); riparian Picea glauca/Populus balsamifera forest with thick moss cover, Peel River, 66°00.36'’N 134°43.19'W, B. Bennett 00-467, 11 July 2000 (DAO). This species is widespread in the Territory (Cody 1996) but is new to the Wind and Peel rivers. Lycopodium clavatum L. var. monostachyon Hook. & Grev., Common Club-moss — YUKON: moist area between cliff and Salix/Populus/Picea forest, Wind River Camp #4, 65°22.89'N 135°26.1'W, B. Bennett 00-204, 5 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 60 kilometers from a site adjacent to the Peel River. To the southwest the nearest site adjacent to the Dempster Highway is about 175 kilometers. Lycopodium selago L., Mountain Club-moss — YUKON: saddle in alpine, upper slope of mountain, Bonnet Plume River east of Fairchild Lake, 64°58'23"N 133°42'15"W, V. Loewen 99-1-2, 11 July 1999 (DAO); steep active solifluxion slope, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-250, 3 July 2000 (DAO); in deep moss in open Picea glauca forest, Wind River, 64°48.46'N 134°41.34'W, B. Bennett 00-831, 3 July 2000 (DAO); steep talus slope, Wind River Camp #4, 65°22.89'N 135°26.1’W, B. Bennett 00-172, 5 July 2000 (DAO). This is a widespread species in the Territory (Cody 1996) but the specimens cited above fill in a blank area in the Wernecke Mountains area. SELAGINELLACEAE Selaginella selaginoides (L.) Link, Spikemoss — YUKON: amongst Parnassia in moss at edge of river, Wind River, McClusky Lake, 64°34.19'N 134°25.77'W, B. Bennett 00-332, 2 July 2000 (DAO); at muddy base of steep active solifluxion slope, Wind River Camp #2, 64°51.83'N 134°38.85'W, B. Bennett 00-161, 4 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory of about 175 kilometers to the _ east and northeast of sites adjacent to the Dempster Highway and just north of latitude 64°N. ISOETACEAE Isoetes echinospora Dur., Bristle-like Quillwort — 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 449 YUKON: on shore of small pond south of Morris Lake, 60°20'N 131°40’W, R. Rosie 2030, 8 Sept. 1999 (DAO) (determined by D. Brunton 2000). This is a rare species in the Yukon Territory (Douglas et al. 1981). The specimen cited above is an extension of the known range in the Territory of about 160 kilometers southwest of a site in the vicinity of Frances Lake where Rosie (1991) found it in 1979. A collection by Porsild and Breitung from Sheldon Lake adjacent to the North Canol Road originally treated as Isoetes braunii by Porsild (1951) and as J. echinospora by Cody (1996) was recently revised to I. maritima by Brunton and Britton (1999). EQUISETACEAE Equisetum fluviatile L., Water Horsetail — YUKON: slough, red rock seep near confluence with the Peel River, Wind River, 65°50’N 135°18'W, B. Bennett 00-452, 8 July 2000 (DAO); valley bottom, palus- trine emergent sedge wetland, near small pond, Bonnet Plume River, 65°02’20"N 134°17'25’W, V. Loewen 99-25-82, 14 July 1999 (DAO). Cody (1996) mapped this species throughout much of the Territory. The specimens cited above are new records for the Wind and Bonnet Plume rivers. Equisetum palustre L., Marsh Horsetail — YUKON: edge of slough, Wind River Camp #1, 64°40.39'N 134°35.96"W. B. Bennett 00-718, 2 July 2000 (B. Bennett Herbarium, photo DAO); base of mountain above Bonnet Plume River, 64°59'40"N 134°03'32"W, V. Loewen 99-8-35, 12 July 1999 (DAO); Larix laricina/Picea mariana/Sphagnum bog, Peel River Camp #10, between Snake and Bonnet Plume rivers, 65°57.61'N 134°25.68'W, B. Bennett 00-696, 12 July 2000 (B. Bennett Herbarium, photo DAO); in sandy lake shallows at west end of Morris Lake, 60°26’N 131°43’W, R. Rosie 2025, 10 Sept. 1999 (DAO); Salix/Betula thickets, Nisutlin Delta, Colwell Bay, 60°13.11'N 132°23'W, B. Bennett 98-656, 28 July 1998 (DAO). The specimens cited above extend the known range about 200 kilometers from sites adjacent to the Dempster Highway (Cody 1996) and 150 kilometers south of a site adjacent to the South Canol Road. Equisetum pratense Ehrh., Meadow Horsetail — YUKON: in young Populus balsamifera riparian for- est on riverbar, Taco Bar, Peel River, 66°00.15'N 134°13'W, B. Bennett 00-398, 13 July 2000 (DAO); Richardson Mts., in mature White Spruce/Balsam Poplar forest on floodplain of Caribou River, 66°15'40"N 135°31'28"W, R. Rosie 2146, 15 July 2000 (DAO); Richardson Mts., river alder, 66°50'22"N 134°56'56’"W, G. Brunner 27-00, July 2000 (DAO); Richardson Mts., alder/rose, 66°52’N 135°50'W, G. Brunner 20-00, July 2000 (DAO). The nearest site of this species to the specimens cited above known to Cody (1996) is about 175 kilometers to the northwest north of the Dempster Highway. Equisetum scirpoides Michx., Dwarf Scouring-rush — YUKON: on steep vegetated slope above river, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-719, 2 July 2000 (B. Bennett Herbarium, photo DAO). This species is found throughout much of the Territory (Cody 1996). The specimen cited above is intermediate between sites between the Hart and Blackstone rivers and a site adjacent to the Snake River (Cody et al. 2001). ASPIDIACEAE Cystopteris fragilis (L.) Bernh., Fragile Fern — YUKON: steep calcareous slopes forming moist caves with talus, east bank of Wind River near confluence of Royal Creek, 65°06.83’N 135°06.43'W, B. Bennett 00-264, 3 July 2000 (DAO); in calcareous talus, Peel River Site #1, 65°58.6’N 134°49.69’'W, B. Bennett 00-405, 10 July 2000 (DAO). The specimens cited above are extensions of the known range to the east of specimens mapped by Cody (1996) of about 75 kilometers. Cystopteris montana (Lam.) Bernh., Mountain Bladder Fern — YUKON: moist rich river bottom with step-moss, open Picea glauca forest on old river terrace, Wind River Camp #6, 65°40.46'N 135°11.76'’W, B. Bennett 00-850, 7 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 150 kilometers to the east. Gymnocarpium jessoense (Koidz.) Koidz. ssp. parvulum Sarvela, Nahanni Oak Fern — YUKON: wooded steep draws with steep active talus shale under Alnus crispa, junction of Wind and Peel rivers, 65°50.48’N 135°18.25'W, B. Bennett 00-813, 8 July 2000 (DAO); in calcareous talus at edge of Picea/Alnus thicket, Peel River Site #1, 65°58.06'N 134°49.69'W, B. Bennett 00-409, 10 July 2000 (DAO). Douglas et al. (1981) (sub. G. robertiana) considered this species rare in the Territory. The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 200 kilometers to the northeast from a site adjacent to the Dempster Highway. CUPRESSACEAE Juniperus communis L. ssp. depressa (Pursh) Franco, Ground Juniper — YUKON: gravel roadside, Alaska Highway 21 km NW of Lake Creek Campsite, 61°58'33.8"”N 140°27'50.3"W, Cody & Cody 37138, 26 July 2000 (DAO). The specimen cited above is an extension of the known distribution of this widespread species in the Yukon Territory of about 100 kilometers northwest of a site adja- cent to the north end of Kluane Lake. Juniperus horizontalis Moench, Creeping Juniper — YUKON: creek valley growing in limestone outwash, Wind River Camp #3, 65°06.83'N 135°06.43'W, B. Bennett 00-150, 3 July 2000 (DAO). The specimen cited above is the northernmost yet found in the Territory. The nearest site mapped by Cody (1996) is adjacent to the Bonnet Plume River about 60 kilometers to the southeast. 450 PINACEAE Picea mariana (Mill.) B.S.P., Black Spruce — YUKON: Alaska Highway, 2.3 km SE of border crossing, 62°36'05.2”N 140°58'04.9"W, Cody & Cody 37217, 26 July 2000 (DAO); hummocky Picea mariana site south of recent fire west of cat trail, upper aufeis site, Wind River, 64°58.67'N 134°47.12’W, B. Bennett 00-307, 4 July 2000 (DAO). The nearest site mapped by Cody (1996) to the first location cited above is about 100 kilometers to the south- east adjacent to the Alaska Highway. The second specimen is from a site in the Wernecke Mountains intermediate between locations south of latitude 64°N and.just south of the Peel River. Pinus contorta Doug]. ex Loud. ssp. latifolia (Engelm.) Critchfield, Lodgepole Pine — YUKON: single tree on east side of road in shallow soil over rock on slope above road among Salix shrubs and Epilobium angustifolium, Dempster Highway Km 72.5, 1.8 km N of Tombstone Mountain Camp- ground, Cody et al. 37023, 22 July 2000 (DAO); small stand of pine on both sides of highway, Alaska Highway Km 1581, 60°48.81’N 136°49.42’W, B. Bennett 00-678, 16 July 2000 (DAO). The first specimen cited above is the northernmost Lodgepole Pine yet known in the Yukon Territory. It is an extension of the known range in the Territory (Cody 1996) of about 175 kilometers northwest of a site adjacent to the Klondike Highway and 160 kilometers west by northwest of a site in the vicinity of Mayo. The second specimen that is the westernmost yet found in the south is about 90 kilo- meters west of the vicinity of Whitehorse. SPARGANIACEAE Sparganium minimum (Hartm.) Fries, Small Bur- reed — YUKON: slough, No Gold Creek Horseshoe Slough, 63°26'N 135°06’W, D. Mossop s.n., 1 July 1999 (B. Bennett Herbarium, photo DAO); Taye Lake, 60°56'N 136°21'W, M. Dennington s.n., 21 July 1982 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The first specimen cited above is about 175 kilo- meters to the north of a site adjacent to the Yukon River and about 200 kilometers southeast of the vicinity of Dawson City mapped by Cody (1996). The second speci- men cited above is from a site about 125 kilometers south- west of the Yukon River site mapped by Cody (1996). Sparganium multipedunculatum (Morong) Rydb., Bur-reed — YUKON: small pond by South McQuesten River near Elsa, 63°56.2'N 135°34.4'W, S. Withers 528, 10 Aug. 1999 (DAO). The specimen cited above is an extension of the known range in the Territory of about 150 kilometers east of a site near the junction of the Klondike and Dempster highways (Cody 1996). POTAMOGETONACEAE Potamogeton friesii Rupr., Flat-stalk Pondweed — YUKON: in small pond near lake, Aishihik Lake, 61°38'N 137°28'W, R. Rosie 2052, 30 July 1998 THE CANADIAN FIELD-NATURALIST py | Vol. 116 (DAO); in shallows of small upland lake, Frances Lake, 61°11'N 129°08'W, R. Rosie 2009, 7 July 1999 (DAO); in shallows of small slough along Frances River, R. Rosie 2011, 5 Aug. 1998 (DAO). The specimens cited above are range extensions from localities in the Territory (Cody 1996): Aishihik Lake about 90 kilometers north of Haines Junction and Frances Lake about 125 kilometers north of Watson Lake. Potamogeton richardsonii (Benn.) Rydb., Richard- son’s Pondweed — YUKON: pond off river, Wind River Site #2, 64°54.2'N 134°41.25'’W, B. Bennett 00-160, 4 July 2000 (DAO). The specimen cited above is from a location intermedi- ate between a site just north of latitude 64°N (Cody 1996) and a site just north of latitude 66°N cited by Cody et al. (2001). SCHEUCHZERIACEAE Scheuchzeria palustris L. ssp. americana (Fern.) Hultén, Scheuchzeria — YUKON: in quaking bog near Morris Lake, 60°25'16”N 131°40'46’W, R. Rosie 2033, 9 Sept. 1999 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is an extension of the known range in the Territory of about 175 kilometers west of a site reported by Cody et al. (1998) in the vicinity of Watson Lake. Triglochin maritimum L., Seaside Arrow-grass — YUKON: muddy slough on riverbar, Wind River, 65°12.49'N 135°13.17'W, B. Bennett 00-136, 5 July 2000 (DAO); boggy tussock tundra near small pond, Wind River Camp #2, 64°51.83'N 134°38.85’'W, B. Bennett 00-815, 4 July 2000 (DAO); in quaking bog near Morris Lake, 60°26'N 131°43'’W, R. Rosie 2034, 10 Sept. 1999 (DAO). The specimens cited above are extensions of the known range in the Territory (Cody 1996) of about 180 kilometers southeast of the Dempster Highway, 150 kilometers south of a site north of latitude 66°N cited by Cody et al. (2001) and intermittent between Whitehorse and Watson Lake. ALISMATACEAE (ALISMACEAE) Sagittaria cuneata Sheld., Arum-leaved Arrowhead — YUKON: < 50 plants seen in shallows, Little Atlin Lake, 60°17.84'’N 133°59.15’W, B. Bennett 00-592, 30 July 2000 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is an extension of the known range in the Territory (Cody 1996) from sites east and northeast adjacent to the Alaska Highway. POACEAE Alopecurus pratensis L., Meadow Foxtail — YUKON: gravel roadside, Alaska Highway, 21 km NW of Lake Creek Campground, 61°58'33.8"”N 140°27'50.3”W, Cody & Cody 37135, 26 July 2000 (DAO); disturbed gravel at roadstop, Alaska Highway 74 km NW of Snag Junction Campground, 62°17'16.4"N .140°46'45.5"W, Cody & Cody 37186, 26 July 2000 (DAO); Mechanic Creek west of Big Creek drainage, 62°20'N 137°19.5'W, S. Withers 212, 21 July 1999 (B. Bennett Herbarium, photo DAO). 2002 Coby, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 451 Cody (1996) knew this introduced species from only three widely separated locations in the Territory. Cody et al. (2001) added two additional sites at Elsa and south of Whitehorse. The specimens cited above extend the known range in the Territory to near the Alaska border adjacent to the Alaska Highway and about 150 kilometers to the east of the highway. Bromus inermis Leyss., Smooth Brome — YUKON: Lake Creek, 61°51'19.6"N 140°09'11.2”W, Cody & Cody 37124, 26 July 2000 (DAO); broad gravel roadside, Alaska Highway, 400 m NW of White River Bridge 61°59'27.4”N 140°33'55.6"W, Cody & Cody 37144, 26 July 2000 (DAO); roadside gravel slope, Alaska Highway, 20 km NW of White River Bridge, 62°08'39.6'N 140°41'05.0"W, Cody & Cody 37160, 26 July 2000 (DAO); Beaver Creek Com- munity, 62°23'11.1”N 140°52'28.3"W, Cody & Cody 37228, 26 July 2000 (DAO). The specimens cited above are a range extension of this introduced species in the Territory of about 200 kilometers on the Alaska Highway to near the Alaska border from a site northwest of Haines Junction (Cody 1996). Bromus pumpellianus Scribn. var. arcticus (Shear) A.E. Porsild, Pumpelly Brome — YUKON: open riverbar in gravels and silt, Illytd Creek confluence with Wind River, 65°30.07'N 135°22.88'W, B. Bennett 00-108, 6 July 2000 (B. Bennett Herbarium, photo DAO); loose sand near top of beach, Taco Bar, Peel River, 66°00.15’N 134°13'W, B. Bennett 00-389, 13 July 2000 (B. Bennett Herbarium, photo DAO). This is a widespread species in the Territory (Cody 1996) but the only specimens between latitude 64° and 66°30'N were adjacent to the Dempster Highway and to the west. Calamagrostis canadensis (Michx.) Beauv. ssp. langsdorfii (Link) Hultén — YUKON: disturbed grav- elly soil of lower slope of ski hill, Mt. Sima Ski Hill, Whitehorse, 60°36'41.2”N 135°02'46.6"W, Cody & Cody 37477, 4 Aug. 2000 (DAO); gravel slope, Alaska Highway, just east of Alaska border line, 62°36'53.8"N 140°59'59.8’W, Cody & Cody 37195, 37196 (DAO); valley bottom near small pond, Bonnet Plume River, 65°02'20"N 134°17'25’W, V. Loewen 99-26-83, 14 July 1999 (DAO). This subspecies that is widespread throughout much of the Territory (Cody 1996) has not previously been found in the vicinities of Whitehorse and the Alaska Highway near the Alaska border, and in the area between latitudes 64°N and 67°N east of the Dempster Highway from a single col- lection northwest of the Bonnet Plume site between the Wind and Hart rivers. Calamagrostis stricta (Timm) Koeler ssp. stricta, Slimstem Reedgrass — YUKON: riverbar, Wind River Camp #6, 65°40.46'’N 135°11.76'W, B. Bennett 00-148, 7 July 2000 (B. Bennett Herbarium, photo DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 100 kilometers east of the Dempster Highway. Elymus calderi Barkworth, Calder’s Wild Rye — YUKON: riverbar, Wind River, 64°24.43'N 134°28.88'W, B. Bennett 00-478, 2 July 2000 (DAO); moderately steep east slope of west-facing cirque, Bonnet Plume River, 64°50’50"N 133°18'37"W, V. Loewen 99-36-102, 16 July 1999 (DAO); lower ridge, rocky slope, Upper Bonnet Plume River Drainage Site #133, 64°35'45"N 132°47'50"W, J. Staniforth 00-088, 8 July 2000 (DAO). Cody (1996) knew this species in the Territory only north to latitude 64°N. Cody et al. (1998) reported a new site adjacent to the Dempster Highway. The specimens cited above are from about 260 kilometers east of the Dempster Highway. Elymus elongatus (Host) Runemark ssp. ponticus (Podp.) Melderis, Tall Wheat Grass (Figure 1) — YUKON: roadside gravel at Yukon/British Columbia border, 24.5 road kilometers south of Carcross, 60°00'00.6’”N 134°39'47.1"W, Cody & Cody 37406, 30 July 2000 (DAO). This invasive plant which is new to the Yukon Territory originated in Turkey and was planted in the prairies as a salt-tolerant forage. It is now known from all the provinces but Newfoundland, Prince Edward Island, New Brunswick and the Northwest Territories. Elymus elongatus ssp. ponti- cus can be separated from E. spicatus as follows: A. Glumes thin, flexible, acute; culms 10—100 cm tall; native species found on dry riverbanks and slopes PMR hs oP ne ot ES a See es ee E. spicatus A. Glumes thick, stiff, obtuse to truncate; culms 70—200 cm tall; introduced, found in disturbed sites such as roadsides...... E. elongatus ssp. ponticus Elymus glaucus Buckl., Western Rye Grass — YUKON: steep stony grassy slope adjacent to road on north side of Yukon River, Whitehorse, 1.1 km N of Grey Mountain cemetery, 60°42'21.0"N 134°59'52.0’"W, Cody & Cody 37422, 31 July 2000 (DAO). Douglas et al. (1981) considered this species rare in the Territory and has since been reported in the extreme south- east (Cody et al. 2000). The specimen cited above is an extension of the known range in the Territory of about 150 kilometers to the east from sites in the Kluane area. Elymus macrourus (Turcz.) Tzvelev, Thick-spike Wild Rye — YUKON: between foot of cliff and road, Canol Road Km 202, 61°53'30"N 132°45'07.5"W, Cody and Cody 36937, 18 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory of about 100 kilometers to the east of a site adjacent to the Yukon River (Cody 1996). Elymus trachycaulus (Link) Gould ex Shinners ssp. glaucus (Pease & Moore) Cody — YUKON: borders of roads in campground by Salix, Alnus and Picea, Pine Valley Campground, Alaska Hwy., 61°51'14.3”N 140°08'58.9"W, Cody & Cody 37111, 26 July 2000 (DAQ). The specimen cited above is an extension of the known range in the Territory of about 75 kilometers to the north- west from a site at the north end of Kluane Lake (Cody 1996). Elymus elongatus © 200) FIGURE 1. Elymus elongatus ssp. ponticus, Tall Wheat Grass (Drawn by Lee Mennell). Elymus trachycaulus (Link) Gould ex Shinners ssp. novae-angliae (Scribner) Tzvelev — YUKON: dis- turbed gravel at roadstop, Alaska Highway 7.4 km NW of Snag Junction Campground, 62°17'16.4”N 140°46'45.5"W, Cody & Cody 37180, 26 July 2000 (DAO) (determined by S. J. Darbyshire). The specimen cited above is an extension of the known range in the Territory of about 175 kilometers northwest of a site at the south end of Kluane Lake (Cody 1996). Festuca baffinensis Polunin, Baffin Fescue — YUKON: alpine tundra, Kotaneelee Range, 60°15’N 124°08'W, R. Rosie 2072, 20 June 1998 (DAO); moist draw in subalpine meadow, Kotaneelee Range, 60°14.31’N 124°07.19'W, B. Bennett 98-710, 15 June 1998 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 400 — kilometers southwest of a site near Ross River. Festuca brachyphylla Schultes & Schultes f., Short- THE CANADIAN FIELD-NATURALIST a ) Vol. 116 leaf Fescue — YUKON: on sandy flats at north end of small lake, Frances Lake, 61°12’N 129°05’W, R. Rosie 2005, 5 July 1999 (DAO). The specimen cited above is a extension of the known range in the Territory (Cody 1996) of about 100 kilometers west of a site just west of longitude 127°W (Cody et al. 1998). Festuca lenensis Drob., Tundra Fescue — YUKON: This mining trench was reclaimed in 1998 using an experimental agronomic seed mix, Division Mountain near Nordenskiold, 61°19’N 136°03.5’W, S. Withers 320, 26 July 1999 (B. Bennett Herbarium, photo DAO); mining trench, Red Ridge, 60°21.5’N 135°04.2'W, S. Withers 109, 14 July 1999 (DAO). Cody (1996) knew this species in the Yukon Territory north of latitude 67°N with a single site in the south in Kluane National Park. The specimens cited above are from about 230 kilometers northeast and 200 kilometers south- east of the Kluane site. Festuca rubra L. sl., Red Fescue — YUKON: roadside gravel, 24.5 road kilometers south of Carcross at the Yukon/BC border, 60°00'00.6"N 134°39'47.1"W, Cody & Cody 37405, 30 July 2000 (DAO); borders of roads in campground by Salix, Alnus and Picea, Pine Valley Campground, Alaska Highway, 61°51'14.3”N 140°08'58.9"W, Cody & Cody 37106, 26 July 2000 (DAO); Alaska Highway 21 km NW of Lake Creek Campground, 61°58'33.8’N 140°27'50.3”W, Cody & Cody 37134, 26 July 2000 (DAO); broad gravel road- side, Alaska Highway, 400 m NW of White River bridge, 61°59'27.4"N 140°33'55.6"W, Cody & Cody 37143, 26 July 2000 (DAO); gravel at rest stop, Alaska Highway 21 km NW of White River bridge, 62°08'51.6"N 140°41'04.5"W, Cody & Cody 37166, 26 July 2000 (DAO). The specimens cited above extend the known range in the Territory (Cody 1996) about 100 kilometers south from a site west of Whitehorse and about 160 kilometers north- west of the north end of Kluane Lake. Festuca trachyphylla (Hackel) Krajina, Hard Fescue — YUKON: borders of roads in campground by Salix, Alnus and Picea, Pine Valley Campground, Alaska Highway, 61°51'14.3”N 140°08'18.9"W, Cody & Cody 37104, 26 July 2000 (DAO). Cody (1996) knew this introduced species only from the vicinity of Dawson City where Calder and Billard collected it in 1949, Festuca vivipara (L.) Sm. ssp. glabra Frederikson, Viviparous Fescue — YUKON: solifluxion plain in valley, Vuntut National Park, 68°25.45’N 138°41.27'W, B. Bennett 00-505, 5 Aug. 2000 (DAO). The specimen cited above is an extension of the known range in the far north of the Territory (Cody 1996) of about 75 kilometers to the south from a site in the British Mountains. Hierochloe hirta (Schrank) Borbas ssp. arctica G. Weim., Common Sweet Grass — YUKON: riverbar, 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 453 Wind River Camp #1, 64°40.39'N 134°35.96’W, B. Bennett 00-354, 2 July 2000 (DAO); open riverbar in gravels, Illytd Creek confluence with Wind River, 65°30.07'N 135°22.88'W, B. Bennett 00-109, 6 July 2000 (DAO); loose sand near top of beach, Taco Bar, Peel River, 66°00.15’N 134°13'W, B. Bennett 00-391, 13 July 2000 (DAO). The specimens cited above are the first known in the Territory from east of the Dempster Highway (Cody 1996), an extension of the known range to the east of about 175 kilometers. Hordeum jubatum L., Foxtail Barley — YUKON: gravel slope, Alaska Highway just east of Alaska border line, 62°36'52.8"”N 140°59'59.8"W, Cody & Cody 37200, 26 July 2000 (DAO); slumping eroding banks, cobbles and silty muck, Wind River, 65°47.45'N 135°13'W, B. Bennett 00-120 (DAO); loose talus at base of steep active scree slope, junc- tion of Wind and Peel rivers, 65°50.48'’N 135°18.25'’W, B. Bennett 00-810, 8 July 2000 (DAO). The specimens cited above are extensions of the known range in the Territory (Cody 1996) of about 100 kilometers to the northwest from a site on the Alaska Highway in the vicinity of the Donjek River and about 99 kilometers east of a site just east of the Hart River. Phalaris arundinacea L., Reed Canary Grass — YUKON: French Gulch, north of Eldorado Creek drainage, 63°53.8’N 139°21.1'W, S. Withers 332, 28 July 1999 (B. Bennett Herbarium, photo DAO); reclaimed mining trench, Mechanic Creek, 62°20’N 137°19.5'W, S. Withers 203, 21 July 1999 (DAO). Mining trenches at these sites were reclaimed in 1995 using an experimental agronomic seed mix that may have included Phalaris arundinacea. The nearest sites to those listed above are about 125 kilometers to the southeast adja- cent to the Klondike Highway and about 250 kilometers to the north adjacent to the Dempster Highway. Douglas et al. (1981) considered this species rare in the Territory. The introduction of races of this species for reclamation may cause genetic contamination of the native race. Phleum pratense L., Timothy — YUKON: broad gravel roadside, Alaska Highway 400 m NW of White River bridge, 61°59'27.4"N 140°33'55.6’W, Cody & Cody 37151, 26 July 2000 (DAO); roadside gravel slope, Alaska Highway 20 km NW of White River Bridge, 62°08'39.6”N 140°41’05.0"W, Cody & Cody 37161, 26 July 2000 (DAO). The specimens cited above extend the known range of this introduced species in the Territory (Cody 1996) about 170 kilometers to the northwest from a site at the south end of Kluane Lake. Poa abbreviata R.Br. ssp. abbreviata, Northern Blue Grass — YUKON: ground shrub/herb/moss-lichen, Bonnet Plume River, 64°40'06"N 133°13'20"W, V. Loewen 99-28-90a, 15 July 1999 (DAO). This is a rare species in the Yukon Territory (Cody 1996). The specimen cited above is an extension of the known range in the Territory of about 320 kilometers east of a site northwest of Dawson City. Poa alpina L., Alpine Blue Grass —— YUKON: wet sand by stream, Koidern River (Edith Creek) behind the Pine Valley Campground, 61°51'19.6”N 140°09'11.2"W, Cody & Cody 37129, 26 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory adjacent to the Alaska Highway of about 100 kilometers from a site near Destruction Bay (Cody 1996). Poa annua L., Annual Blue Grass — YUKON: sandy gravel cleared area, Top of the World Highway Km 29.2, Cody & Cody 37037, 22 July 2000 (DAO). This is an uncommon introduced species in the Territory (Cody 1996). It was known to Hultén (1968) from the vicinity of Dawson City but has not been found there recently. Cody et al. (1998) reported it from the Beaver River area in the southeast. It is a new introduction along the Top of the World Highway. Poa arctica R.Br. ssp. arctica, Arctic Blue Grass — YUKON: moderately steep talus slope, Bonnet Plume River, 64°44'37"N 133°24'30’"W, V. Loewen 99-38- 119, 16 July 1999 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 100 kilometers northeast of a site north of Mayo. Poa glauca Vahl, Glaucous Blue Grass — YUKON: borders of roadsides in campground by Salix, Alnus and Picea, Pine Valley Campground, Alaska High- way, 61°59'14.3"N 140°08'58.9"W, Cody & Cody 37109, 26 July 2000 (DAO); broad gravel roadside, Alaska Highway 400 m NW of White River bridge, 61°59'27.4'N 140°33'55.6"W, Cody & Cody 37142, 26 July 2000 (DAO). This is a widespread species in the Territory (Cody 1996) but has not previously been collected adjacent to the Alaska Highway northwest of Kluane Lake although it is known from the mountains to the west. Poa pratensis L. ssp. pratensis, Kentucky Blue Grass — YUKON: Koidern River (Edith Creek) behind the Pine Valley Campground, 61°51'19.6"N 140°09'11.2"W, Cody & Cody 37126, 26 July 2000 (DAO). This is a widespread introduced species in the Territory, particularly south of latitude 64°N (Cody 1996). The speci- men cited above is from a new site adjacent to the Alaska Highway between Kluane Lake and the Alaska border. Stipa hymenoides Roem. & Schult., Indian Rice Grass — YUKON: disturbed sandy area across road from snow dump, above Whitehorse on north side of Yukon River, 60°42'44.4"N 135°01'26.6"W, Cody & Cody 36916, 17 July 2000 (DAO). Cody et al. (2001) reported this species new to the Yukon Territory on the basis of a specimen collected by Bennett on a sandy eroding slope beside the hydro dam at the Schwatka Lake Dam in the Yukon River at Whitehorse. The specimen cited above is from a site about 2 kilometers to the north of the site by the hydro dam. Trisetum sibiricum Rupr. ssp. sibiricum, Siberian Flase Oat — YUKON: lower part of steep rocky slope 454 above road, Tatchun/Frenchman Road Km 28, 20 km E of Klondike Highway, 62°14'46.5"N 135°56'33.7"W, Cody & Cody 36993, 21 July 2000 (DAO). This rare amphi-beringian species was previously known from only two sites in the Territory (Cody 1996): Porcupine River and Coffee Creek on the Yukon River. Trisetum spicatum (L.) Richt., Spike Trisetum — YUKON: riverbar, Wind River Camp #4, 65°22.89'N 135°26.1'W, B. Bennett 00-371, 5 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 75 kilometers to the south from a site adjacent to the Bonnet Plume River just south of the Peel River. CYPERACEAE Carex aquatilis Wahlenb. ssp. aquatilis, Water Sedge — YUKON: palustrine emergent sedge wet- land, valley bottom, Bonnet Plume River, 65°02'20"N 134°17'25"W, V. Loewen 99-25-81, 14 July 1999 (DAO); headwaters of Bonnet Plume River, Mackenzie Mountains, 64°20’N 132°35'W, J. Meikle s.n., 5 Aug. 1997 (DAO); silty mud on edge of riverbank, Wind River Camp #1, 64°30.39'N 134°35.96'’W, B. Bennett 00-762, 2 July 2000 (DAO); emergent from lake, McClusky Lake, Wind River, 64°34.19'N 134°25.77'W, B. Bennett 00-287, 2 July 2000 (DAO). The specimens cited above are extensions of the known range of this species of about 75 kilometers to the northeast and about 200 kilometers to the east of a site in the Wernecke Mountains west of longitude 135°W (Cody 1996). Carex concinna R.Br., Low Northern Sedge — YUKON: older Picea glauca/Populus balsamifera for- est beside creek in shady mossy area, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00- 731, 2 July 2000 (DAO); occasional in moist Picea glauca woods, Wind River Camp #4, 65°22.89'N 135°26.1'W, B. Bennett 00-261, 5 July 2000 (DAO); open Picea glauca forest on old river terrace, Wind River Camp #6, 65°40.46’N 135°11.76'W, B. Bennett 00-863, 7 July 2000 (DAO); riparian spruce rorest,. Peel River ‘Camp #10, 65°57.61' N 134°25.68'W, B. Bennett 00-684, 12 July 2000 (DAO). The specimens cited above are the first known between latitudes 64°N and 66°N east of the Dempster Highway. Carex gynocrates Wormskj., Yellow Bog Sedge — YUKON: on steep vegetated slope above river, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-716, 2 July 2000 (DAO); moderately steep east slope of west-facing cirque, Bonnet Plume River, 64°50'50"N 133°18'37"W, V. Loewen 99-36- 105, 16 July 1999 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 175 kilometers northeast of a site in the vicinity of Mayo. Carex lasiocarpa Ehrh. ssp. americana (Fern.) THE CANADIAN FIELD-NATURALIST ee 7 Voli Hultén, Wooly-fruit Sedge — YUKON: with Carex utriculata along small creek in wetland, Frances Lake, 61°09’N 129°07'W, R. Rosie 2001, 11 July 1999 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimens cited above are an extension of about 125 kilometers north of a site adjacent to Watson Lake reported by Cody et al. (1998). Carex lugens Holm, Spruce Muskeg Sedge — YUKON: woodland white spruce/shrub stand near Bonnet Plume River, 64°59'48”N 134°04'10’W, V. Loewen 99-41-134, 16 July 1999 (DAO); severely mounded lower slope, tussock tundra-Carex/ Eriophorum, Upper Bonnet Plume River Drainage, 64°19'23"N 131°35'9"W, J. Staniforth 00-123, 00- 125, July 2000 (DAO). The first specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 150 kilometers to the northeast from a site in the vicinity of Mayo. The second specimen is from a site about 200 kilo- meters east of the Mayo area. Carex misandra R.Br., Short-leaved Sedge — YUKON: cirque lake shore, Bonnet Plume River, 64°50'51"N 133°18'33"W, V. Loewen 99-37-114, 16 July 1999 (DAO); steep northeast-facing alpine slope, Bonnet Plume River, 64°54'36"N 133°34'20"W, V. Loewen 99-10-46, 12 July 1999 (DAO); open dolomite delta, Wind River, 64°48.46'N 134°41.34'W, B. Bennett 00-254, 3 July 2000 (DAO). The specimens listed above are an extension of the known range in the Territory (Cody 1996) of about 140 kilometers to the east from a site in the Wernecke Mountains. Carex nardina Fries, Spikenard Sedge — YUKON: moderately steep east slope of west-facing cirque, Bonnet Plume River, 64°50’50”N 133°18'37’W, V. Loewen 99-36-103, 16 July 2000 (DAO); steep active solifluxion slope across river from camp, Wind River Camp #1, 64°40.38'N 134°35.96’W, B. Bennett 00-1044, 3 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 150 kilometers northeast of a site in the vicinity of Mayo. Carex pellita Willd. — YUKON: large patch at west- ern edge at high water mark, sandy soil, Ear Lake, Whitehorse, 60°40.81'N 135°02.54'W, B. Bennett 00-908, 5 Sept. 2000 (DAO) (determined by P.M. Catling & W.J. Cody). This is a rare species in the Territory (Cody et al. 2000). The specimen upon which this was based was collected adjacent to the Yukon River below Rink Rapids. The speci- men cited above is the second collection known and is from a site about 200 kilometers southeast of Rink Rapids. Carex podocarpa R.Br., Graceful Mountain Sedge — YUKON: exposed windy alpine knoll, Mt. Granger, 60°34.11'N 135°09.38'W, B. Bennett 99- 104, 10 July 1999 (DAO); moderate south-facing slope, Bonnet Plume River, 64°40'06"N 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 455 133°13'20"W, V. Loewen 99-28-89, 15 July 1999 (DAO); steep southeast-facing alpine slope, Bonnet Plume River, 64°53'13"N 133°49'45"W, V. Loewen 99-16-58, 13 July 1999 (DAO); moderately steep east-facing slope east of Fairchild Lake, Bonnet Plume River, 64°58'03”N 133°41'12”W, V. Loewen 99-4-17, 11 July 1999 (DAO). The first specimen cited above is new to the area adja- cent to Whitehorse (Cody 1996) and the remaining speci- mens are an extension of the known range in the Territory of about 100 kilometers north of a site just north of latitude 64°N. Carex rupestris All., Curly or Rock Sedge — YUKON: steep northeast-facing alpine slope, Bonnet Plume River, 64°54'44"N 133°33'54’W, V. Loewen 99-11-52, 12 July 1999 (DAO); undulating lower slope, Dryas outwash floodplain, Upper Bonnet Plume Drainage Site #110, 64°32'10"N 132°19'40"W, J. Staniforth 00-054, 6 July 2000 (DAO); steep active solifluxion slope across river from camp, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-176, 3 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 275 kilometers east of a site adjacent to the Blackstone River. Carex saxatilis L., Russet Sedge —- YUKON: sparse low white spruce/Dryas/cottongrass/moss, Bonnet Plume River, 64°59’40"N 134°03’32"W, V. Loewen 99-8-32, 99-8-37, 12 July 1999 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 150 kilometers to the northeast from a site in the vicinity of Mayo. Carex scirpoidea Michx., Single-spike Sedge — YUKON: riverbar, shrubby Salix, Wind River Camp #1, 64°44.43'N 134°28.88'W, B. Bennett 00-210, 2 July 2000 (DAO); riverbar, Wind River, raven nest site, 65°12.49’N 135°13.17'W, B. Bennett 00-1049, 5 July 2000 (B. Bennett Herbarium); Betula neoalaskana/Salix scouleriana/S. bebbiana/Alnus crispa upland esker in gravelly soil, Wind River Camp #6, 65°40.46’N 135°11.76'W, B. Bennett 00- 864, 7 July 2000 (DAO); steep southeast-facing upper slope, Bonnet Plume River, V. Loewen 99-29- 85, 99-29-84, 15 July 1999 (DAO); moderately steep east slope of west-facing cirque, Bonnet Plume River, 64°50'50"N 133°18'37"W, V. Loewen 99-36- 104, 16 July 1999 (DAO); alpine ridge between two cirques, 64°51'31”N 133°34’50"W, V. Loewen 99-9- 44a, 99-9-44b, 12 July 1999 (DAO). The specimens cited above are intermediate between sites mapped by Cody (1996) on the Upper Bonnet Plume River and adjacent to the Hart River. Carex spectabilis Dewey, Northern Show Sedge — YUKON: in alpine meadow, Morris Lake, 60°23'N 131°33'W, R. Rosie 2021, 11 Sept. 1999 (DAO). This species was considered rare in the Territory by Douglas et al. (1981) who reported it from the Peel River and southeastern Kluane National Park and cited the map in Hultén (1968). That map however did not include a Peel River site and was not included in Cody (1996). The Morris Lake site is an extension of about 340 kilometers east of the Kluane area. Carex tenuiflora Wahlenb., Sparse-leaved Sedge — YUKON: in hummock spruce with Betula glandulosa with open ponds of standing water, Wolf Lake, 60°42.6'N 131°44.08'W, B. Bennett 99-331, 11 Aug. 1999 (DAO). The specimen cited above is an extension of the known range in the south of the Territory (Cody 1996) of about 125 kilometers northwest of a site west of Watson Lake adjacent to the Alaska Highway. Carex vaginata Tausch., Sheathed Sedge — YUKON: alpine ridge between two cirques, Bonnet Plume River, 64°54'31”"N 133°34’'50”W, V. Loewen 99-9- 43, 12 July 1999 (DAO); moist rivulet on east-facing mountain, Wind River Camp #1, 64°30.39'N 134°35.96'W, B. Bennett 00-722, 2 July 2000 (DAO); on hummocky Picea/Salix near river, Wind River, McClusky Lake, 64°34.19'N 134°25.77'W, B. Bennett 00-330, 2 July 2000 (DAO); moist area below cliff with springs, Wind River Camp #4, 65°22.89'N 135°26.1'W, B. Bennett 00-842, 5 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 170 kilometers northwest from a site adjacent to the Upper Bonnet Plume River. Eleocharis uniglumis (Link) Schult., Creeping Spike-rush — YUKON: graminoid/sedge meadows at edge of lake out to mud flats, south end of Stevens Lake, 61°41'40.14"N 137°30'12.63”"W, Staniforth & Rosie 98-167, 28 Aug. 1998 (DAO). Douglas et al. (1981) considered this species rare in the Territory. This specimen cited above is intermediate between sites near Haines Junction and Carmacks (Cody 1996). Eriophorum angustifolium Honckn., Narrow-leaved Cotton-grass — YUKON: boggy tussock tundra near small pond, Wind River Camp #2, 64°51.83'N 134°38.85'W, B. Bennett 00-816, 4 July 2000 (DAO); cirque lake shore, Bonnet Plume River, 64°50'51”N 133°18'33”"W, V. Loewen 99-37-107, 16 July 1999 (DAO); moderate southeast-facing slope of creek valley on east-facing mountain slope, Bonnet Plume River, 64°53'27"N 133°48’01"W, V. Loewen 99-19-72, 13 July 1999 (DAO); Bonnet Plume Lake, 64°20'N 132°W, W.H. Butler & N. Olsen 22, 11-27 July 1966 (CAN). The specimens cited above of this widespread species in the Territory (Cody 1996) are the first known from between latitudes 64°N and 66°N east of the Dempster Highway. Eriophorum vaginatum L., Sheathed Cotton-grass — YUKON: base of mountain above Bonnet Plume River, 64°59’40"N 134°03'32"W, V. Loewen 99-8- 36, 12 July 1999 (DAO); tussock meadow near creek in Bonnet Plume River Valley, 64°54'49"N 456 133°44'05’W, V. Loewen 99-39-125, 16 July 1999 (DAO); severely mounded lower slope, tussock tun- dra, Upper Bonnet Plume River Drainage Site #140, 64°18'23"N 131°35'9"W, J. Staniforth 00-124, July 2000 (DAO). The specimens cited above are an extension of the known range of this widespread species in the Territory (Cody 1996) of about 250 kilometers to the southeast from a site south of the Peel River between the Wind and Hart rivers. ARACEAE Calla palustris L., Wild Calla — YUKON: No Gold Creek, Horseshoe Slough, 63°26'N 135°06’W, D. Mossop s.n., 1 July 1999 (DAO). This is a frequent species in the vicinity of the Porcupine River but is rare in the Territory south of latitude 67°N. The specimen cited above is an extension of about 50 kilometers to the southeast from the vicinity of Mayo. JUNCAECAE Juncus alpinoarticulatus Chaix in Vill ssp. ameri- canus (Farwell) Hamet-Ahti, Alpine Rush — YUKON: moist sandy gravel shore of Long Lake, Whitehorse, 60°44'38.8"N 135°02'47.2"W, Cody & Cody 37461, 3 Aug. 2000 (DAO); silty seep at edge of river, Peel River Site #1, 65°58.06'’N 134°49.69'W, B. Bennett 00-404, 10 July 2000 (DAO). The nearest sites to the first specimen cited above known to Cody (1996) were southwest of Johnson’s Crossing about 100 kilometers to the east and in Kluane National Park about 100 kilometers to the west. The second specimen is intermediate between a site in the vicinity of Mayo and the Porcupine River. Juncus balticus Willd. var. alaskanus (Hultén) A.E. Porsild, Baltic Rush — YUKON: riverbar, Wind River Camp #1, 64°34.43’N 134°28.88'W, B. Bennett 00- 473, 2 July 2000 (DAO); silty mud on edge of lake, Wind River Camp #2, 64°51.83’N 134°38.85'W, B. Bennett 00-827, 4 July 2000 (DAO); muddy slough on riverbar, Wind River, 65°12.49’N 135°13.17'W, B. Bennett 00-457, 5 July 2000 (B. Bennett Herbar- ium, photo DAO). The specimens cited above are intermediate between sites mapped by Cody (1996) adjacent to the Dempster Highway and upper Bonnet Plume River. Juncus castaneus Smith ssp. castaneus, Chestnut Rush — YUKON: in hummocky Picea/Salix forest near river, McClusky Lake, Wind River, 64°34.19'N 134°25.77'W, B. Bennett 00-337, 2 July 2000 (DAO). The specimen cited above is intermediate between sites mapped by Cody (1996) adjacent to the Dempster Highway, upper Bonnet Plume River and just south of latitude 64°N. Juncus castaneus Smith ssp. leucochlamys (Zinz.) Hultén — YUKON: riverbar at base of cliff, Wind River, 65°12.49'N 135°13.17'W, B. Bennett 00-367, 5 July 2000 (DAO). The nearest site known to Cody (1996) to the specimen cited above is in the vicinity of Mayo about 175 kilometers to the south. THE CANADIAN FIELD-NATURALIST Vol. 116 Juncus triglumis L. ssp. albescens (Lange) Hultén, Whitish Rush — YUKON: riverbar, Wind River near confluence of Royal Creek, 65°06.83’N 134°47.12'W, B. Bennett 00-196, 4 July 2000 (DAO); muddy slough on riverbar, Wind River, 65°12.49'N 135°13.17'W, B. Bennett 00-137, 5 July 2000 (DAO). The specimens cited above are intermediate between sites adjacent to the upper Bonnet Plume River and between the Blackstone and Hart rivers mapped by Cody (1996). Luzula arctica Blytt ssp. arctica, Arctic Woodrush — YUKON: open meadow near outfitter’s camp, Wind River, McClusky Lake, 64°34.19’N 134°25.77'W, B. Bennett 00-242, 2 July 2000 (DAO). The specimen cited above is intermediate between sites at about 64°N and 66°N mapped by Cody (1996). Luzula confusa Lindebl., Confused Woodrush — YUKON: moderate south-facing slope, Bonnet Plume River, 64°40'06"N 133°13'20"W, V. Loewen 99-28- 91, 15 July 1999 (DAO); moderately steep talus slope, Bonnet Plume River, 64°44'37"N 133°24'30"W, V. Loewen 99-38-123, 99-38-120, 16 July 1999 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 100 kilometers to the northeast from a site just south of 64°N. Luzula parviflora (Ehrh.) Desv. ssp. parviflora, Small-flowered Woodrush — YUKON: moist dis- turbed area by river’s edge, McClusky Lake, Wind River, 64°34.19’N 134°25.77'W, B. Bennett 00-284, 2 July 2000 (DAO); undulating lower slope, Upper Bonnet Plume River Drainage Site #104, 64°26'15’N 132°15'47’"W, J. Staniforth 00-027, 5 July 2000 (DAO); strongly mounded undulating alpine valley lower slope, Upper Bonnet Plume River Drainage Site #154, 64°28'19"N 132°19'21"W, J. Staniforth 00-154, 12 July 2000 (DAO); undulating upper slope, moist hummocky tundra, Upper Bonnet Plume River Drainage Site #108, 64°28’05"N 132°04'15"W, J. Staniforth 00- 037, 5 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 100 kilometers north-northeast of a site south of latitude 64°N and 140 kilometers northeast of the same site. Luzula rufescens Fisch. & Mey., Rusty Woodrush — YUKON: in willow shrub/birch/Festuca altaica stand on Frances River floodplain, 61°11'N 129°13’30"W, R. Rosie 2006, 19 June 1999 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 350 kilometers east of a site north of Aishihik Lake. Luzula spicata (L.) DC., Spiked Woodrush — YUKON: moderately steep east-facing alpine slope east of Fairchild Lake, Bonnet Plume River, 64°58'09"N 133°41'56”"W, V. Loewen 99-2-10, 11 July 1999 (DAO). 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 457 The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 140 kilometers northeast of sites south of latitude 64°N and is the northern- most yet found. LILIACEAE Allium schoenoprasum L. ssp. sibiricum (L.) Celak, Wild Onion — YUKON: open riverbar in gravels, Wind River, 65°30.07'N 135°22.88'W, B. Bennett 00-107, 6 July 2000 (B. Bennett Herbarium, photo DAO); riverbar on tributary stream in gravels, Wind River, 65°46.36’N 135°10.71'W, B. Bennett 00-124, 8 July 2000 (DAO); riverbar, Peel River Camp #8, 65°56.03'N 134°58.84'W, B. Bennett 00-767, 9 July 2000 (DAO). . The specimens cited above are intermediate between sites mapped by Cody (1996) south of latitude 64°N and north of latitude 67°N. Tofieldia coccinea Richards., Northern False Asphodel — YUKON: talus slope near river, Deception Mtn., Wind River, 65°36.03'’N 135°28.41'W, B. Bennett 00-424, 7 July 2000 (DAO); shallow soil over limestone, Snake River, 65°24'N 133°24’W, J. Meikle 99-008, 20 July 1999 (Yukon Renewable Resources, photo DAQ); steep solifluxion slope, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-174, 3 July 2000 (DAO); steep northeast- facing slope, Bonnet Plume River, 64°54'44”N 133°33'54"W, V. Loewen 99-11-50, 12 July 1999 (DAO). The specimens cited above are intermediate between sites mapped by Cody (1996) and a site adjacent to the Snake River (Cody et al. 2001). ORCHIDACEAE Amerorchis rotundifolia (Banks) Hultén, Round- leaved Orchis — YUKON: moist Picea glauca/Alnus thicket, riparian with seeps, Peel River Site #1, 65°58.06'N 134°49.69'W, B. Bennett 00-402, 10 July 2000 (DAO); riparian Picea glauca/Populus balsamifera/Alnus incana forest, Deception Mountain, Wind River, 65°36.03'N 135°28.41'W, B. Bennett 00-273, 7 July 2000 (DAO); wet area between talus and Picea glauca/Alnus incana woods, Wind River Camp #4, 65°22.89'N 135°26.1'W, B. Bennett 00-259, 5 July 2000 (DAO); steep moist draw in Populus tremuloides/Picea glauca/Ledum groenlandicum stand in subalpine, Wind River Camp #3, 65°06.83'’N 135°06.43'W, B. Bennett 00- 151, 3 July 2000 (DAO). The specimens cited above extend the known distribu- tion of this species in the Territory (Cody 1996) south into the Wernecke Mts. about 110 kilometers from two sites just north of latitude 66°N. Calypso bulbosa (L.) Oakes, Fairy-slipper — YUKON: open Picea glauca forest on old river terrace, Wind River Camp #6, 65°40.46’N 135°11.76'W, B. Bennett 00-253, 7 July 2000 (B. Bennett Herbarium, photo DAO). The specimen cited above is an extension of the known range in the Territory of about 250 kilometers north of the vicinity of Mayo and about 290 kilometers northeast of the vicinity of Dawson City (Cody 1996). Coeloglossum viride (L.) Hartm. ssp. bracteatum (Muhl.) Hultén, Bracted Green Orchid — YUKON: deciduous shrub-Salix/Betula/ ericaceous shrubs, Donjek Burn, 61°30'N 139°36'’W, C.E. Kennedy s.n., 26 June 1985 (Yukon Renewable Resources, photo DAO); riverbar, Wind River Camp #3, 65°06.83'N 134°47.12'W, B. Bennett 00-192, 4 July 2000 (DAO). Douglas et al. (1981) considered this species rare in the Territory and was known to Cody (1996) from only two widely separated localities: adjacent to Atlin Road in the extreme south and adjacent to the Bonnet Plume River far to the north. Cody et al. (2001) reported a specimen from the Snake River to the east of the Bonnet Plume River. The Wind River collection is from west of the Bonnet Plume River site. The Donjek Burn collection is from a site about 350 kilometers northwest of the Atlin Road site. Corallorhiza trifida Chat., Striped Coralroot — YUKON: Picea mariana/Betula papyrifera/Sphagnum bog on top of bluffs, junction of Wind and Peel rivers, 65°50.48'N 135°18.25'W, B. Bennett 00-112, 9 July 2000 (DAO); riverbar, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-355, 2 July 2000 (DAO); in hummocky Picea/Salix forest near river, McClusky Lake, Wind River, 64°34.19'N 134°25.77'W, B. Bennett 00-375, 2 July 2000 (DAO). This is a widespread species in the Territory (Cody 1996) but was previously unknown east of the Dempster Highway between latitudes 64°N and 66°N. Cypripedium guttatum Sw., Spotted Lady’s-slipper — YUKON: moist areas well up on slope with Larix laricina, Deception Mountain, Wind River, 65°36.03'N 135°28.41'W, B. Bennett 00-426, 7 July 2000 (DAO); moist glade in Picea glauca forest above open dolomite delta, Wind River, 64°48.46'N 134°41.34'W, B. Bennett 00-254, 3 July 2000 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimens cited above extend the known range in the Territory south into the Wernecke Mountains about 120 kilometers from a site near the Peel River (Cody 1996). Cypripedium parviflorum Salisb. var. makasin (Farwell) Sheviak (Sheviak 1993) (C. calceolus L. ssp. parviflorum (of recent authors), Small Yellow Lady’s-slipper — YUKON: moist wooded slope growing in talus, Wind River Camp #3, 65°06.83'N 134°47.12'W, B. Bennett 00-214, 4 July 2000 (DAO); moist areas well up on slope with Larix lar- icina, Deception Mountain, Wind River, 65°36.03'N 135°28.41'W, B. Bennett 00-270, 7 July 2000 (B. Bennett Herbarium, photo DAQ). Douglas et al. (1981) considered this species rare in the Territory. Cody (1996) knew it from only four localities: vicinity of Faro, two adjacent to the Bonnet Plume River and adjacent to the Alaska border at 66°N. Cody et al. 458 (2000) extended the known range to the northeast of the Alaska border site. Goodyera repens (L.) R.Br., Dwarf Rattlesnake- plantain — YUKON: wet moss under spruce, Alaska Highway, 2.3 km SE of border crossing, 62°36'05.2"N 140°58'04.9"W, Cody & Cody 37214, 26 July 2000 (DAO). The nearest site of this orchid species to the locality in the Territory is about 75 kilometers due south adjacent to the Alaska border (Cody 1996). Platanthera obtusata (Pursh) Lindl., Northern Bog Orchid — YUKON: Riparian Picea glauca/Populus balsamifera/Alnus incana forest, Deception Mountain, Wind River, 65°36.03'’N 135°28.41'W, B. Bennett 00- 425, 7 July 2000 (DAO); at base of steep talus slope beside muddy seep, Wind River Camp #4, 65°22.89'N 135°26.1'W, B. Bennett 00-856, 5 July 2000 (DAO); Picea glauca riparian forest, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-361, 2 July 2000 (DAO); undulating spruce/shrub/ lowshrub/forb/moss area, Upper Bonnet Plume River Drainage near Site #105, 64°26'6"N 132°15'28’W, J. Staniforth 00-011, 5 July 2000. The specimens cited above are from sites up to about 100 kilometers west and northwest and southeast from a site adjacent to the Snake River reported by Cody et al. (2001), the only other known location between latitudes 64°N and 66°N east of the Dempster Highway. Spiranthes romanzoffiana Cham. & Schlecht., Hooded Ladies’ -tresses — YUKON: wet moss, Aishihik Lake, Km 42 on Aishihik road from Alaska Highway, 61°11'38.3”N 136°59'54.9"W, Cody & Cody 37312, 29 July 2000 (DAO). The specimen cited above is from a site intermediate between the vicinity of Whitehorse about 100 kilometers to the southeast and a site adjacent to the Alaska Highway about 200 kilometers to the northwest (Cody 1996). SALICACEAE Populus balsamifera L. ssp. balsamifera, Balsam Poplar — YUKON: older Picea glauca/Populus bal- samifera forest beside creek, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-727, 2 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 125 kilometers to the northeast from a site in the vicinity of Mayo. Populus tremuloides Michx., Trembling Aspen — YUKON: open stand in steep draw in subalpine, Wind River Camp #3, 65°06.83’N 135°06.43' W, B. Bennett 00-149, 3 July 2000 (DAO); steep open slope, Deception Mountain, Wind River, 65°36.03'N 135°28.41'W, B. Bennett 00-274, 7 July 2000 (DAO). The specimens cited above are intermediate between sites mapped by Cody (1996) between the vicinity of Mayo and latitude 66°N. Salix arbusculoides Anderss., Northern Bush Willow — YUKON: moist rich river bottom with step-moss, THE CANADIAN FIELD-NATURALIST Vol. 116 open Picea glauca forest on old river terrace, Wind River Camp #6, 65°40.46'N 135°11.76'W, B. Bennett 00-851, 7 July 2000 (DAO); riparian forest, Wind River, 65°12'49"N 135°13'17"W, B. Bennett 00-139, 5 July 2000 (DAO); loose talus at base of steep active shale scree slope, junction of Wind and Peel rivers, 65°50.48'’N 135°18.25'’W, B. Bennett 00-804, 8 July 2000 (DAO); older Populus balsam- ifera/Salix forest beside creek between Snake and Bonnet Plume rivers, Peel River Camp #9, 65°58.13’N 134°48.61'W, B. Bennett 00-793, 00- 791, 10 July 2000 (DAO); edge of river in older Salix alaxensis/S. pulchra/Calamagrostis canadensis meadow, Peel River Camp #10, 65°57'61"N 134°25'68"W, B. Bennett 00-713, 12 July 2000 (DAO); woodland white spruce/shrub stand near Bonnet Plume River, 64°59'48"N 134°04'10’W, V. Loewen 99-4]-132, 17 July 1999 (DAO) (deter- mined by G. Argus). The specimens cited above are an extension of the known range in the Territory of about 150 kilometers from sites adjacent to the Dempster Highway. Salix barrattiana Hook., Barratt’s Willow — YUKON: base of mountain above Bonnet Plume River, 64°59'40"N 134°03'32"W, V. Loewen 99-8-40, 12 July 1999 (DAO); wetland undulating valley floor sedge meadow, Upper Bonnet Plume River Drainage Site #142, 64°22'47"N 132°07'52’W, J. Staniforth 00- 133, 10 July 2000 (DAO); shrub meadow valley bot- tom, Upper Bonnet Plume River Site #146, 64°22'26"N 132°46'40"W, J. Staniforth 00-143, 10 July 2000 (DAO); undulating upper slope, moist hum- mocky tundra, Upper Bonnet Plume River Drainage Site #108, 64°28’05"N 132°04'11"W, J. Staniforth 00- 038, 5 July 2000 (DAO); valley floor, moist shrub/graminoid/moss, Upper Bonnet Plume River Drainage Site #114, 64°24’51”"N 132°23’'W, J. Staniforth 00-065, 6 July 2000 (DAO); riverbar, Wind River, Camp #1, 64°34'43"N 134°28'88"W, B. Bennett 00-267, 2 July 2000 (DAO) (determined by G. Argus). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 250 kilometers east of sites between the Hart and Blackstone rivers. Salix myrtillifolia Anderss., Bilberry Willow — YUKON: marshy area near river, McClusky Lake, Wind River, 64°34.19’N 134°25.77'W, B. Bennett 00-241, 2 July 2000 (DAO); older Picea glauca/Populus balsamifera forest beside creek, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-749, 2 July 2000 (DAO); base of moun- tain above Bonnet Plume River, 64°59’ 40"N 134°03'32"W, V. Loewen 99-8-38, 12 July 1999 (DAO) (determined by G. Argus). The specimens cited above are an extension of the known range in the Territory of about 75 kilometers to the north of a site just north of latitude 64°N (Cody 1996). 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 459 Salix planifolia Pursh ssp. pulchra (Cham.) Argus — YUKON: moderately steep N-facing slope, Bonnet Plume River, 65°01'48"N 134°18'55”W, V. Loewen 99-23-77, 14 July 1999 (DAO); tussock meadow near creek in Bonnet Plume River valley, 64°54'49"N 133°44’05"W, V. Loewen 99-39-124, 16 July 1999 (DAO); open meadow and shrubby area near outfitter’s camp, McClusky Lake, Wind River, 64°34.19’N 134°25.77'W, B. Bennett 00- 312, 00-234, 2 July 2000 (DAO) (determined by G. Argus). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 100 kilometers to the north of a site just north of latitude 64°N. Salix pseudomonticola Bebb (S. monticola sensu Cody 1996), Mountain Willow — YUKON: edge of river in older Salix alaxensis/S. pseudomonticola/S. pulchra/Calamagrostis meadow, Peel River Camp #10 between Snake and Bonnet Plume rivers, 65°57.61'N 134°25.68'W, B. Bennett 00-871, 12 July 2000 (DAO) (determined by G. Argus). The specimen cited above is the northernmost yet found in the Territory (Cody 1996). It is from about 275 kilome- ters north of a site in the vicinity of Mayo. Salix rotundifolia Trautv. ssp. dodgeana (Rydb.) Argus — YUKON: moderately steep southwest-facing slope, Bonnet Plume River, 64°41'30’"N 133°05'15"W, V. Loewen 99-30-87, 15 July 1999 (DAO) (determined by G. Argus). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 100 kilometers to the southeast. BETULACEAE Betula glandulosa Michx., Ground Birch — YUKON: on steep vegetated slope above river, Wind River Camp #1, 64°40.39'N 134°35.96’W, B. Bennett 00- 755, 2 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 120 kilometers northeast from a site north of Mayo. Betula neoalaskana Sarg., Alaska Birch — YUKON: edge of Picea between low Salix, Alaska Hwy. 2.3 km SE of border crossing, 62°36'05”N 140°58'04.9"W, Cody & Cody 37219, 26 July 2000 (DAO). This is a widespread species in the Territory north to just beyond latitude 68°N. The specimen cited above is from a location about 60 kilometers northwest of a site adjacent to the Alaska Highway (Cody 1996). SANTALACEAE Geocaulon lividum (Richards.) Fern., Northern Comandra — YUKON: steep vegetated slope above river, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-701, 2 July 2000 (DAO).; hummocky Picea mariana site south of recent fire west of cat trail, Wind River, 64°58.67'N 134°47.12'W, B. Bennett 00-306, 4 July 2000 (DAO). The specimens cited above are from sites about 75 kilo- meters south of a location adjacent to the Bonnet Plume River cited by Cody et al. (2000). POLYGONACEAE Polygonum buxiforme Small, Eastern Knotweed — YUKON: gravel disturbed area, Alaska Highway at Destruction Bay, 61°15’07.6"N 138°48'19.8"W, Cody & Cody 37069, 25 July 2000 (DAO); gravel beside Alaska Highway, Beaver Creek Community, 62°23'11.1"N 140°52'28.3”"W, Cody & Cody 37224, 26 July 2000 (DAO). This native weedy species which is found as far north adjacent to the Dempster Highway to north of latitude 66°N (Cody 1996) is now known northwest of the vicinity of Haines Junction adjacent to the Alaska Highway near the Alaska border. Polygonum viviparum L., Alpine Bistort — YUKON: amongst Salix on riverbar, Wind River, 65°12.49'N 135°13.17'W, B. Bennett 00-458, 5 July 2000 (DAO); on steep vegetated slope above river, Wind River Camp #1, 69°40.39'N 134°35.96'W, B. Bennett 00-738, 2 July 2000 (DAO). The specimens cited above are intermediate between a site adjacent to the Snake River to the east (Cody et al. 2001) and a site adjacent to the Hart River to the west. Rheum rhaponticum L., Rhubarb — YUKON: grow- ing wild in town area, Destruction Bay, 61°15'10.2”N 138°40'00.5"W, Cody & Cody 37233, 27 July 2000 (DAO). Cody et al. (2001) reported Rhubarb growing wild at Silver City on the east side of the south end of Kluane Lake. Rumex crispus L., Curled Dock — YUKON: waste ground along roadside, Alaska Highway, 16.5 km W of Klondike Highway, 60°51'34.2"N 135°28'24.8"W, Cody & Cody 37052, 24 July 2000 (DAO); gravel waste area, Haines Junction, 60°45'06.7"N 137°30'28.7”W, Cody & Cody 37062, 25 July 2000 (DAO); gravel slope, Alaska Highway, just east of Alaska border line, 62°36'52.8"N 140°59'59.8"W, Cody & Cody 37208, 37209, 26 July 2000 (DAO). Cody (1996) knew this species that was introduced from Europe only from the vicinity of Dawson City. CHENOPODIACEAE Chenopodium album L., Lamb's-quarters — YUKON: disturbed outfitter camp, Wind River Camp #2, 64°45.05'N 134°38.75'W, B. Bennett 00-185, 3 July 2000 (DAO). The specimen cited above of this introduced species is an extension of the known range of about 150 kilometers to the northeast from a site adjacent to Mayo (Cody 1996). Salicornia europaea L., Slender Glasswort — YUKON: dry narrow meadow between rock outcrops, Albert Rock’s grazing lease, south of Deep Creek rock, Lake Laberge, 61°03'29.1'N 135°11'44.4"W, P. Smith 00-865, 10 Aug. 2000 (DAQ). Douglas et al. (1981) considered this species rare in the Territory. Cody (1996) knew it from only two localities ee 460 adjacent to the Alaska Highway. The specimen cited above is an extension of the known range of about 60 kilometers to the northeast. CARYOPHYLLACEAE Arenaria longipedunculata Hultén, Low Sandwort — YUKON: alpine tundra, Aishihik L. area, 61°57'N 37°50'W, R. Rosie 2042, 15 Aug. 1999 (DAO). Cody (1996) knew this species from widely scattered areas in the Territory. The specimen cited above is from a locality about 140 kilometers north of a site in Kluane National Park. Minuartia biflora (L.) Schinz. & Thell., Mountain Sandwort — YUKON: alpine ridge between two cirques, Bonnet Plume River, 64°54'31"N 133°34'50”"W, V. Loewen 99-9-43a, 12 July 1999 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 160 kilometers to the northeast of sites south of latitude 64°N. Minuartia rossii (R.Br.) Graebn., Ross’ Stitchwort — YUKON: alpine barren on ridge of mountain sum- mit west of camp, Vuntut National Park, 68°27.52'N 138°46.02'’W, B. Bennett 00-519, 7 Aug. 2000 (DAO); vicinity of Snowdrift Camp, Vuntut National Park, 68°21.4’N 139°13.1'W, P. Caswell PPC-2000-Y-102, 20 June 2000 (B. Bennett Herbarium, photo DAO). Cody (1996) knew this species in the southwest of the Territory and adjacent to the Arctic coast. The specimens cited above are an extension of about 70 kilometers to the southwest into the mountains. Moehringia lateriflora (L.) Fenzl, Blunt-leaved Sand- wort — YUKON: midslope shrub zone, Upper Bonnet Plume River Drainage Site #150, 64°30'7”"N 133°01'36"W, J. Staniforth 00-146, 11 July 2000 (DAO); loose talus at top of steep active shale slope, junction of Wind and Peel rivers, 65°50.48'N 135°18.25'W, B. Bennett 00-802, 8 July 2000 (DAO); talus at base of cliff, Taco Bar, Peel River, 65°55’N 134°15'W, B. Bennett 00-386, 12 July 2000 (DAO); alder/rose, Richardson Mts., 66°44'51”N 135°24'40"W, G. Brunner 24C-00, July 2000 (DAO). The specimens cited above are intermediate between sites adjacent to the Dempster Highway and Porcupine River (Cody 1996) and a site adjacent to the Snake River (Cody et al. 2001). Silene acaulis L. ssp. subacaulescens (F. N. Wil- liams) Hultén, Moss Campion — YUKON: on steep vegetated slope above river, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-753, 2 July 2000 (B. Bennett Herbarium, photo DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 120 kilometers northeast from a site north of Mayo. Stellaria longipes Goldie, Long-stalked Starwort — YUKON: moderately steep north-facing slope, Bonnet Plume River, 65°02'03”N 134°19’09”"W, V. Loewen 99-24-78, 14 July 1999 (DAO); open meadow near THE CANADIAN FIELD-NATURALIST Vol. 116 outfitter’s camp, Wind River, McClusky Lake, 64°34.19’N 134°25.77'W, B. Bennett 00-311, 00- 350, 2 July 2000 (DAO). The specimens cited above are intermediate between a site just east of the Hart River (Cody 1996) and a site adja- cent to the Snake River (Cody et al. 2001). Wilhelmsia physodes (Fisch.) McNeill — YUKON: edge of river, McClusky Lake, Wind River, 64°34.19'N 134°25.77'W, B. Bennett 00-335, 2 July 2000 (DAO); riverbar at base of cliff, Wind River, 65°12.49'N 135°13.17'W, B. Bennett 00-364, 5 July 2000 (DAO); open riverbar in gravels, Illytd Creek confluence with Wind River, 65°30.07'N 135°22.88'W, B. Bennett 00-104, 6 July 2000 (B. Bennett Herbarium, photo DAO); riverbar of tribu- tary stream in gravels, Wind River, 65°46.36'’N 135°10.71'W, B. Bennett 00-126, 8 July 2000 (B. Bennett Herbarium, photo DAO); riverbar in cob- bles, Peel River Camp #10 between Snake and Bonnet Plume rivers, 65°57.61'N 134°25.68'W, B. Bennett 00-705, 12 July 2000 (DAO). This species is frequent adjacent to the Dempster Highway between latitudes 64° and 66° (Cody 1996) but with the exception of a single collection from the upper Bonnet Plume River was unknown to the east. NYMPHAEACEAE Nuphar polysepalum Engelm., Yellow Water-lily — YUKON: shallow water near lakeshore, Enger Lakes, 62°14'23.7"N 140°40'52.8"W, Cody & Cody 37176, 26 July 2000 (DAO). The specimen cited above is an extension to the known range in the Territory of about 150 kilometers to the vicini- ty of the Alaska Highway in the southwest from a site adja- cent to the junction of the Yukon and Stewart rivers (Cody 1996). RANUNCULACEAE Ranunculus cooleyae Vasey & Rose, Plume Anemone (Figure 2). Martha Louise Black (1944) in her book of Yukon Wild Flowers included a photograph of this Buttercup and provided the following information: “We found these flowers for the first time in Yukon last summer. Mr. Porsild of the Dominion botanical staff, tells me this plant is entirely new to the Yukon. The name Plume Anemoneae comes from the beauti- fully plume like arrangement of the stamens. The flower is ivory white, the stamens lemon yellow, the pistils a deeper yellow. Skelly, a prospector friend at Carcross, first brought our attention to this newcom- er. I must here pay a tribute to that friend who has often helped us in our search for local wild flowers”. This species was not included in the Flora of the Yukon Territory (Cody 1996) and unfortunately has not yet been rediscovered. _ Ranunculus cymbalaria Pursh, Northern Seaside Buttercup — YUKON: open disturbed area, south end of Canol Road, 60°29'26.0”N 133°17'42.0"W, Cody & Cody 36927, 18 July 2000 (DAO). 2002 Coby, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 461 FIGURE 2. Ranunculus cooleyae (Drawn by Valerie Fulford). The specimen cited above is an extension of the known range in the Territory of about 115 kilometers to the east from a site in the vicinity of Whitehorse (Cody 1996). Ranunculus nivalis L., Snow Buttercup — YUKON: along wet trail dominated by sedge, Larsen Lake, 60°07'00"N 125°33’40’W, V. Loewen 99-2, 12 June 1999 (DAO); alpine meadow near camp, Beaver- crow Ridge, 60°12.9'N 124°35.79'W, B. Bennett 98- 647, 16 Aug. 1998 (B. Bennett Herbarium). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 320 kilometers southeast of a site in the vicinity of Frances Lake. BRASSICACEAE (CRUCIFERAE) Arabis columbiana Macoun, Elegant Rockcress — YUKON: broken rocky slope between highway and Windy Arm Tagish Lake, 1.3 km north of B.C. bor- der, 60°00'25.9"N 134°38'40.03"W, Cody & Cody 37414, 30 July 2000 (DAO); Quill Creek, Kluane National Park, 60°40'N 137°22’W, P. Caswell PPC- 2000-Y-196, 17 July 2000 (Yukon Renewable Resources, photo DAO) (determined by G. A. Mulligan). The first specimen of this rare species in the Territory cited above is only the third yet known. The nearest site to those cited above is in the vicinity of Carmacks (Cody 1996) about 175 kilometers northeast of the Kluane National site. Arabis holboellii Hornem. var. secunda (Howell) Jepson, Holboell’s Rockcress — YUKON: disturbed gravelly soil at foot of ski hill, Mt. Sima Ski Area, Whitehorse, 60°36'41.2”N 135°02'46.6"W, Cody & Cody 37471, 4 Aug. 2000 (DAO); top of slope above Long Lake, 3.8 km on Wickstrom/Long Lake road from Hospital Rd. intersection, 60°44’ 38.8"N 135°02'47.2"W, Cody & Cody 37455A, 3 Aug. 2000 (DAO) (determined by G. A. Mulligan). The specimens cited above are an extension of the known range of this variety in the Territory of about 100 kilometers southeast from the nearest known site adjacent to the Klondike Highway (Cody 1996). Arabis nuttallii Robins., Nuttall’s Rockcress — YUKON: mossy meadow adjacent to Coal River Springs, 61°03’31"N 127°23'42’W, J. Staniforth s.n., 9 June 1996 (determined by G. A. Mulligan); old experimental farm (Warden Hg.) NW of Haines Junction, 60°46'N 137°35'W, P. Caswell PPC- 2000-Y-228, 14 June 2000 (DAO); Horse Trail 1 km NW of old experimental farm, 60°46’N 137°37'W, P. Caswell PPC-2000-Y-236, 10 June 2000 (DAO). This is a rare species in the Territory which Cody (1996) knew from only two localities. Three additional sites were added by Cody et al. (2001). The specimens cited above are an extension of about 450 kilometers east of Whitehorse. Arabis pinetorum Tidestrom — YUKON: gravel road- side, Tagish Road 14 km W of Jakes Corner, 60°19'05.4"N 134°10'08.6"W, Cody & Cody 37389B, 30 July 2000 (DAO) (determined by G. A. Mulligan). The specimen cited above is an extension of the known range in the Territory of about 100 kilometers southeast of a site northeast of Whitehorse (Cody 1996). Brassica campestris L. (B. rapa sensu Cody 1996), Bird Rape — YUKON: disturbed farm area, Mayo, 63°35.26'N 135°51.83'W, B. Bennett 00-380, 2 July 2000 (B. Bennett Herbarium, photo DAO). The specimen cited above is an extension of the known range of this introduced species in the Territory (Cody 1996) of about 150 kilometers southeast from a site near the south end of the Dempster Highway. Capsella bursa-pastoris (L.) Medic, Shepherd’s-purse — YUKON: gravel at rest stop, Alaska Highway, 21 km NW of White River Bridge, 62°08'51.6"N 140°41'04.5"W, Cody & Cody 37168B, 26 July 2000 (DAO); disturbed gravel at roadstop, Alaska Highway 7.4 km NW of Snag Junction Campground, 62°17'16.4"N 140°46'45.5"W, Cody & Cody 37184, 26 July 2000 (DAO); disturbed outfitter camp, Wind River Camp #2, 134°38.75'N 64°45.05'W, B. Bennett 00-187, 3 July 2000 (DAO) (determined by G. A. Mulligan). The specimens cited above are extensions of the known range of this introduced species in the Territory (Cody 1996) adjacent to the Alaska Highway of about 225 kilo- meters northwest of Haines Junction and about 130 kilome- ters northeast of the vicinity of Mayo. Cardamine bellidifolia L., Alpine Bitter-cress — YUKON: moderately steep talus slope, Bonnet Plume River, 64°44'37"N 133°24'30’W, V. Loewen 99-38- 122, 16 July 1999 (DAO) (determined by G. A. Mulligan). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 50 kilometers to the east from a site adjacent to the Wind River. A = 462 Cardamine digitata Richards., Richardson's Rock- cress — YUKON: moist area below cliff with springs, Wind River Camp #4, 65°22.89’N 135°26.1'W, B. Bennett 00-840, 5 July 2000 (DAO); exposed rocky alpine slope, with seepages, Upper Blackstone River at Horn Claims, 64°27.1'N 138°38.2'W, S. Withers SW00-64, 14 July 2000 (DAO). The first specimen cited above of this amphi-beringian species is an extension of the known range in the Territory (Cody 1996) of about 150 kilometers east of sites adjacent to the Dempster Highway. The second specimen is the southernmost yet found in the Territory and is from about 40 kilometers south of the nearest Dempster Highway site. Cardamine pratensis L., Cuckoo Bitter-cress — YUKON: marshy area near lake, McClusky Lake, Wind River, 64°34.19’N 134°25.77'W, B. Bennett 00-236, 2 July 2000 (B. Bennett Herbarium, photo DAO); moist area below cliff with springs, Wind River Camp #4, 65°22.89'N 135°26.1'W, B. Bennett 00-838, 5 July 2000 (B. Bennett Herbarium, photo DAO) (determined by G. A. Mulligan). The specimens cited above are intermediate -between sites adjacent to the Dempster Highway and upper Bonnet Plume River (Cody 1996). Descurainia incisa (Engelm. ex A. Gray) Britton var. incisa, Tansy Mustard — YUKON: roadside gravel slope, Alaska Highway, 20 km NW of White River bridge, 62°08'39.6"N 140°41'05.0"W, Cody & Cody 37159, 26 July 2000 (DAO); disturbed gravel at road- stop, Alaska Highway, 7.4 km NW of Snag Junction Campground, 62°17'16.4"N 140°46'45.5’"W, Cody & Cody 37182, 37183, 26 July 2000 (DAO); gravel slope, Alaska Highway just east of Alaska border line, 62°36'53.8"N 140°59'59.8"W, Cody & Cody 37197, 26 July 2000 (DAO) (determined by G. A. Mulligan). Cody et al. (2001) reported this species as new to the Territory from three widely separated localities. The speci- mens cited above are an extension of the known range of about 450 kilometers northwest of a site in the vicinity of Crag Lake on the Tagish Road. Draba alpina L., Alpine Draba — YuKoNn: alpine, mud, shale moderate south-facing slope, V. Loewen 99-28-94, 15 July 1999 (DAO) (determined by G. A. Mulligan). The nearest site in the Territory known to Cody (1996) to the specimen cited above is adjacent to the Dempster Highway about 225 kilometers to the west. Draba borealis DC., Northern Draba — YUKON: alpine tundra, Kotaneelee Range, 60°15'N 124°08'W, R. Rosie 2068, 20 June 1998 (DAO); along ridge, Beavercrow Ridge, 60°13.629'N 124°32.28'W, B. Bennett 98-651, 15 Aug. 1998 (DAO) (determined by G. A. Mulligan). The specimens cited above are an extension of the known range in the Territory of about 180 kilometers to the southeast of a site west of longitude 127° reported by Cody et al. (1998). Draba stenoloba Ledeb., Alaska Whitlow-grass — YUKON: moist meadow, Coal River area, THE CANADIAN FIELD-NATURALIST Vol( iG 60°51'33”N 127°40'49"W, J. Staniforth s.n., 6-9 June 1996, (DAO); vicinity of Snowdrift Camp, Vuntut National Park, 68°21.4’N 139°13.1'W, P. Caswell PPC-2000-Y-051, 26 June 2000 (DAO) (determined by G. A. Mulligan). Cody (1996) knew this species in the southern part of the Territory north to about 63°N and then disjunct to the western part of the British Mountains. The first specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 280 kilometers to the south- east from a site adjacent to the North Canol Road. The sec- ond specimen is from a site about 130 kilometers southeast of the British Mountains site. Erysimum cheiranthoides L., Wormseed Mustard — gravelly disturbed roadside near tramway, Alaska Highway, Km 1890, 62°02'23”N 140°36'55’W, B. Bennett 97-215, 3 June 1997 (DAO); gravel roadside, Alaska Highway, 17 km NW of White River bridge, 62°07'24.9"N 140°30'35.3"W, Cody & Cody 37154, 26 July 2000 (DAO) (determined by G. A. Mulligan). The specimens cited above are from locations about 180 kilometers northwest of a site at the south end of Kluane Lake and west about 170 kilometers of a site adjacent to a tributary of the Kluane River mapped by Cody (1996). Erysimum cheiri (L.) Crantz, Common Wallflower (Figure 3) — YUKON: between Robert Service Way and old railway in grass adjacent to Picea and Populus about 1.3 km from Alaska Highway, Whitehorse, 60°42'14.4’N 135°03'10.8"W, Cody & Cody 37439, 1 Aug. 2000 (DAO) (determined by G. A. Mulligan). This species is a new introduction into the Yukon Territory. It is an infrequent garden escape on southeastern Vancouver Island, British Columbia (Douglas et al. 1998) which was introduced from Europe. Erysimum cheiri can be separated from E. angustatum as follows: A. Stem leaves linear, crowded, canescent from dense malpighiaceous hairs; caudex usually multicipital; plants < 20 cm tall; petals bright yellow’... 9. aes dee eee E. angustatum A. Stem leaves mostly lanceolate to oblanceolate, more or less entire, pubescent with malpighiaceous hairs; caudex with one or a few stems; plants usually more than 20 cm tall; petals orange-yellow . .... 5 ..e: 52.0 eee E. cheiri Erysimum inconspicuum (S. Wats.) MacMill., Shy Wallflower (Figure 4) — YUKON: gravel soil by track below steep rocky slope, Tagish Rd. Km 38, 60°15'09.2"N 134°29'07.3"W, Cody & Cody 37398, 30 July 2000 (DAO) (determined by G. A. Mulligan). On the basis of the revision of specimens from the Yukon Territory and Continental Northwest Territories for- merly identified as Erysimum inconspicuum to E. coarcta- tum, Cody et al. (2000) removed E. inconspicuum from the flora of the Territory. On the basis of the specimen cited above, E. inconspicuum should now be placed back in the flora of the Yukon (Cody 1996) and added to the list of rare species in the Territory (Douglas et al. 1981). Halimolobus mollis (Hook.) Rollins, Soft Hali- molobus — YUKON: roadside gravel slope, Alaska 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 463 Erysimum cheiri © 2001 FIGURE 3. Erysimum cheiri, Common Wallflower, (Drawn by Lee Mennell). Highway 20 km NW of White River bridge, 62°08'39.6"N 140°41'05.0"W, Cody & Cody 37158, 26 July 2000 (DAO) (determined by G. A. Mulligan). The specimen cited above is an extension of the known distribution in the Territory of about 190 kilometers north- west of a site between Haines Junction and Kluane Lake (Cody 1996). Lepidium bourgeauanum Thell., Bourgeau’s Pepper- grass — YUKON: gravel cleared area, Faro, 62°13'59.1"N 133°20'57.3”W, Cody & Cody 36975, 20 July 2000 (DAO); gravel slope, Alaska Highway just east of Alaska border line, 62°36'52.8"N 140°59'59.8"W, Cody & Cody 37211, 26 July 2000 (DAO) (determined by G. A. Mulligan). The Faro specimen cited above is an extension of the known range in the Territory of about 175 kilometers north from Johnsons Crossing and the Alaska Highway specimen is an extension of the known range adjacent to the highway of about 60 kilometers from south of latitude 62°N (Cody 1996). Lepidium ramosissimum A. Nelson, Branched Pepper-grass — YUKON: roadside gravel, Burwash aA SH RY 8a//4 WNP RY LY ra am Erysimum Inconspic wan © 200) FIGURE 4. Erysimum inconspicuum, Shy Wallflower (Drawn by Lee Mennell). Landing, Alaska Highway, 61°21'17.5"N 139°00'04.6"W, Cody & Cody 37093, 25 July 2000 (DAO); old experimental farm NW of Haines Junction, 60°46'N 137°35'’W, P. Caswell PPC- 2000-Y-404, 17 July 2000 (DAO) (determined by G. A. Mulligan). Previously known sites of this introduced species in the Territory were in the vicinity of Tagish, and adjacent to the Klondike Highway (Cody 1996) and adjacent to French- man Lake (Cody et al. 2001). The specimens cited above are the first known from adjacent to the Alaska Highway. Lesquerella arctica (Wormskj.) S. Wats. ssp. calderi (Mulligan & Porsild) Hultén, Arctic Bladderpod — YUKON: steep loose eroding vegetated hillside above river, Wind River, 64°56.67'N 134°47.12'W, B. Bennett 00-297, 4 July 2000 (B. Bennett Herbarium, photo DAQ); talus slope near river, Deception Mountain, Wind River, 65°36.03'N 135°28.41'W, B. Bennett 00-276, 7 July 2000 (DAQ); riverbar, Peel River Camp #8, 65°56'N 134°58.84'W, B. Bennett 00-778, 9 July 2000 (DAQ); cobbly silty riverbar, 464 Peel River Camp #9, 65°58.13’N 134°48.61'W, B. Bennett 00-795, 10 July 2000 (DAO). The specimens cited above are the easternmost yet found in the Territory (Cody 1996). Parrya arctica R.Br., Arctic False Wallflower — YUKON: alpine mud shale moderate south-facing slope, Bonnet Plume River, 64°40'06”N 133°13'20"W, V. Loewen 99-28-93, 15 July 1999 (DAO) (determined by G. A. Mulligan). Cody (1996) knew this species that is an endemic of the Arctic Archipelago only from Herschel Island about 600 kilometers northwest of this Bonnet Plume River site. Parrya nudicaulis (L.) Regel, Northern Parrya — riverbar, Wind River, 64°34.43’N 134°28.88'W, B. Bennett 00-480, 2 July 2000 (DAO); on steep vege- tated slope above river, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-739, 2 July 2000 (B. Bennett Herbarium, photo DAO); high alpine in loose talus, Wind River, 64°48.46'’N 134°41.34’W, B. Bennett 00-447, 3 July 2000 (B. Bennett Herbarium, photo DAO); steep northeast- facing alpine slope, Bonnet Plume River, 64°54'36"N 133°34'20"W, V. Loewen 99-10-48, 12 July 1999 (DAO); woodland white spruce/ shrub stand near Bonnet Plume River, 64°59'50"N 134°04'17’W, V. Loewen 99-40-126, 17 July 1999 (DAO); convex lower ridge, dry well drained rocky slope, Upper Bonnet Plume River Drainage Site #132, 64°31'19"N 132°51'32"W, J. Staniforth 00-094, 7 July 2000 (DAO) (determined by G. A. Mulligan). The specimens cited above extend the known range in the Territory (Cody 1996) about 130 kilometers to the east from a site just west of longitude 135°W. Rorippa barbareifolia (DC.) Kitagawa, Hoary Yellow Cress — YUKON: gravel at rest stop, Alaska Highway, 21 km NW of White River bridge, 62°09'51.6"N 140°41'04.5"W, Cody & Cody 37164, 26 July 2000 (DAO); slope below rest area, Alaska Highway 18 km SE of border crossing, 62°29'57.9"N 140°51'48.4"W, Cody & Cody 37222, 26 July 2000 (DAO) (determined by G. A. Mulligan). Previous collections of this amphi-beringian species in the Territory were all from north of latitude 63°N (Cody 1996). The nearest site to the specimens cited above is about 150 kilometers to the northeast. Rorippa palustris (L.) Besser ssp. palustris, Marsh Yellow Cress — YUKON: gravel at the Dry Creek rest stop, Alaska Highway, 21 km NW of White River bridge, 62°08'51.6"N 140°41'04.5"W, Cody & Cody 37168A, 26 July 2000 (DAO) (determined by G. A. Mulligan). The specimen cited above is an extension of about 150 kilometers northwest of sites adjacent to Kluane Lake (Cody 1996). Sisymbrium altissimum L., Tumble Mustard — YUKON: gravel at the Dry Creek rest stop, Alaska Highway, 21 km NW of White River bridge, THE CANADIAN FIELD-NATURALIST Vol. 116 62°08'51.6"N 140°41'04.5"W, Cody & Cody 37170, 26 July 2000 (DAO) (determined by G. A. Mulligan). This introduced species was previously known in the Territory only from the vicinities of Dawson City and Whitehorse (Cody 1996). Thlaspi arvense L., Penny Cress — YUKON: open meadow, McClusky Lake, Wind River, 64°34.19’N 134°25.77'W, B. Bennett 00-283, 2 July 2000 (DAO). The specimen cited above extends the known range of this introduced species in the Territory about 100 kilome- ters northeast of a site in Keno (Cody et al. 2001). It was rare at McClusky Lake and not observed elsewhere adja- cent to the Wind and Peel rivers. CRASSULACEAE Crassula aquatica (L.) Schoenl., (Tillaea aquatica L.), Pigmyweed (Figure 5) — YUKON: on shore along the road to town, Nares Lake, Carcross, 60°10'N 134°42'W, Ceska & Goward 11765, 6 July 1982 (DAO); shore of river, N side of bridge across Tagish River, Carcross, 60°10'N 134°42'W, Ceska & Goward 11932, 8 July 1982 (DAO). This species is new to the Flora of the Yukon Territory Crassula aquatica ©2001 FIGURE 5. Crassula aquatica, Pygmyweed (Drawn by Lee Mennell). 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 465 and should be added to the list of rare species (Douglas et al. 1981). Elsewhere in the Canadian north it is only known from the vicinity of Yellowknife (Cody 1954). The genus Crassula can be separated from Rhodiola and Sedum as fol- lows: A. Plants terrestrial perennials; flowers 4 or 5 parted; stamens usually 10 B. Flowers dioecious, 4-parted............ Rhodiola Moeriowers petiect, J-parted .. 2.0... 8. Sedum A. Plants of vernal pools and shallow water, annuals; flowers usually 4-parted; REST 2S ee a ee eee Crassula SAXIFRAGACEAE Boykinia richardsonii (Hook.) A. Gray, Richardson's Boykinia — YUKON: growing in moist crag in lime- stone, Jasper Canyon, Snake River, 65°24’N 133°24'N, J. Meikle 99-028, 99-029, 20 July 1999 (Yukon Renewable Resources, photos DAO). The specimens cited above extend the known range in the Territory about 80 kilometers north of a site adjacent to the Bonnet Plume River (Cody 1996). Ribes hudsonianum Richards., Northern Black Currant — YUKON: Larix laricina/Picea mariana, Carex media, Eriophorum vaginatum marsh, Peel River Camp #10 between Snake and Bonnet Plume rivers, 65°57.61'N 134°25.68'W, B. Bennett 00-701, 12 July 2000 (DAO). The specimen listed above is an extension of the known range in the Territory (Cody 1996) of about 200 kilometers to the southeast from a site east of the Porcupine River. Saxifraga lyallii Engler. ssp. hultenii (Calder & Savile) Calder & Taylor, Red-stemmed Saxifrage — YUKON: moderately steep north-facing slope about mid-way up mountain, Bonnet Plume River, 64°51'10"N 133°27'41"W, V. Loewen 99-5-21, 11 July 1999 (DAO). The specimen cited above is the northernmost yet found in the Territory (Cody 1996). It is from a site about 110 kilometers northwest of a site adjacent to the upper Bonnet Plume River. Saxifraga nivalis L., Alpine Saxifrage — YUKON: alpine tundra, Kotaneelee Range, 60°15'N 124°08’W, R. Rosie 2075, 20 June 1998 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 300 kilometers southeast of a site adjacent to the junction of the Campbell Highway and Nahanni Road. Saxifraga serpyllifolia Pursh, Thyme-leaved Saxi- frage — YUKON: steep northeast-facing slope, Bonnet Plume River, 65°01'03”N 134°18'03’"W, V. Loewen 99-21-74, 14 July 1999 (DAO); moderately steep talus slope, Bonnet Plume River, 64°44'37"N 133°24'30"W, V. Loewen 99-38-131, 16 July 1999 (DAO). The specimens cited above are the easternmost yet found in the Territory between latitudes 62°N and 67°N (Cody 1996). ROSACEAE Amelanchier alnifolia (Nutt.) Nutt., Saskatoon — YUKON: dry steep slope below cliffs above coal mine shaft, Tantalus Butte, Carmacks, 62°07.23'N 136°15.23'W, B. Bennett 00-050, 30 May 2000 (DAO). The specimen cited above is intermediate between sites mapped by Cody (1996) adjacent to the southern Klondike Highway and Mayo. Geum aleppicum Jacq. ssp. strictum (Ait.) Clausen, Yellow Avens — YUKON: Koidern River, 61°51'19.6’"N 140°09'11.2"W, Cody & Cody 37125, 26 July 2000 (DAO); gravel beside Alaska Highway, Beaver Creek Community, 62°23'11.1"N 140°52'28.3"W, Cody & Cody 37225, 26 July 2000 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimens cited above extend the known range in the Territory about 175 kilometers adjacent to the Alaska Highway northwest from a site adjacent to southern Kluane Lake. Geum macrophyllum Willd. ssp. perincisum (Rydb.) Hultén, Large-leaved Avens — YUKON: edge of river in older Salix alaxensis/S. pseudomontisola/S. pul- chra/Calamagrostis canadensis meadow, Peel River Camp #10, 65°57.61'N 134°25.68’W, B. Bennett 00- 706, 12 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 190 kilometers northeast of a site adjacent to the Dempster Highway. This species was not observed adjacent to the Wind River and the site above was the only place where it was seen on the Peel River. Potentilla anserina L., Common Silverweed — YUKON: gravel roadside and ditch, Alaska Highway, 1.5 km north of Beaver Creek bridge, 62°22'36.3"N 140°52'35.6"W, Cody & Cody 37192, 26 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 180 kilometers to the northwest from a site near the south end of Kluane Lake. Potentilla litoralis Rydb., Prairie Cinquefoil — YUKON: gravel slope, Alaska Highway just east of the Alaska border line, 62°36'52.8"N 140°59'59.8"W, Cody & Cody 37213, 26 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 75 kilometers to the northwest from a site south of latitude 62°N adjacent to the Alaska Highway. Potentilla norvegica L., Norwegian Cinquefoil — YUKON: gravel slope, Alaska Highway, just east of Alaska border line, 62°36'52.8”N 140°59'59.8"W, Cody & Cody 37204, 26 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 60 kilometers to the northwest from a site south of latitude 62°N. Potentilla palustris (L.) Scop., Marsh Cinquefoil — YUKON: riverbar, Wind River, 65°12.49'N 135°13.17'W, B. Bennett 00-366, 5 July 2000 (DAQ). The specimen cited above is from the only known locali- nt Sooner Of oh ee hy 466 ty between latitudes 64°N and 66°N, east of the Dempster Highway (Cody 1996). Rubus arcticus L. ssp. acaulis (Michx.) Focke, Nagoonberry — YUKON: in hummocky Picea/Salix forest near river, McClusky Lake, Wind River, 64°34.19’N 134°25.77'W, B. Bennett 00-351, 2 July 2000 (DAO); Larix laricina/Picea mariana/ Sphagnam marsh, Peel River, Camp #10, between Snake and Bonnet Plume rivers, 65°57.61'N 134°25.68'W, B. Bennett 00-697, 12 July 2000 (DAO); in rich moist herbaceous meadow adjacent to fork of Dog Creek, 68°29.14'N 138°50.35’W, B. Bennett 00-1107, 6 Aug. 2000 (DAO). The first two specimens cited above are an extension of the known range in the Territory (Cody 1996) between lati- tudes 64°N and 66°N east of the Dempster Highway. The third specimen is the northernmost yet found in the Territory, a range extension of about 60 kilometers north- east of a site just north of latitude 68°N. Rubus idaeus L. s.1., Wild Red Raspberry — YUKON: riparian forest in talus below cliff, Wind River, 65°12.49'N 135°13.17'W, B. Bennett 00-134, 5 July 2000 (DAO); riverbar near more established willows, Peel River Camp #8, 65°56.03’N 134°58.84’W, B. Bennett 00-775, 9 July 2000 (DAO); Richardson Mts., 66°44'51”N 135°24'40"W, G. Brunner 24A-00, July 2000 (Yukon Renewable Resources, photo DAO). The specimens cited above extend the known distribu- tion in the Territory (Cody 1996) about 150 kilometers east of the Dempster Highway. Sanguisorba officinalis L., Great Burnet — YUKON: on open well washed riverbar, only five plants seen and two others at site downstream, Peel River, 65°55.97'N 135°02.99'W, B. Bennett 00-381, 9 July 2000 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 150 kilometers northeast of a site adjacent to the Dempster Highway reported by Cody et al. (1998). Spiraea beauverdiana Schneid., Steven's Meadow- sweet — YUKON: boggy area with hummocks, Peel River, 66°00.36'N 134°43.19'W, B. Bennett 00-469, 11 July 2000 (B. Bennett Herbarium, photo DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 50 kilometers east of a site west of longitude 135°W. To the south the nearest site is adjacent to the upper Bonnet Plume River. FABACEAE (LEGUMINOSAE) Astragalus alpinus L., Alpine Milk-vetch — YUKON: riverbar, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-358, 2 July 2000 (DAQ); open riverbar in gravels, Wind River conflu- ence with Illytd Creek, 65°30.7'N 135°22.88'W, B. Bennett 00-438, 6 July 2000 (DAO). The specimens cited above are the only records from between latitudes 64°N and 66°N and between longitude 131°30'W and the Dempster Highway (Cody 1996). Astragalus australis (L.) Lam., Indian Milk-vetch — THE CANADIAN FIELD-NATURALIST Vol. 116 YUKON: riverbar with shrubby Salix, Wind River Outfitter’s Camp, 64°34.43'N 134°88'W, B. Bennett 00-479, 2 July 2000 (DAO); steep active solifluxion slope, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-178, 3 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 180 kilometers east of sites adjacent to the Dempster Highway. Astragalus bodinii Sheldon, Bodin’s Milk-vetch — YUKON: uncommon in gravels of creek beds, Dog Creek Camp, Vuntut National Park, 68°27.53'N 138°44'47"W, B. Bennett 00-547, 8 Aug. 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 200 kilometers northwest of a site at about 67°N. Astragalus williamsii Rydb., William’s Milk-vetch — YUKON: clear area adjacent to highway, Alaska Highway 4.0 km S of White River bridge, 61°58'56.3"N 140°28'36.0"W, Cody & Cody 37230, 27 July 2000 (DAO). The specimen cited above is an extension of about 100 kilometers northwest of a site at the north end of Kluane Lake (Cody 1996). Melilotus alba Desc., White Sweet-clover — YUKON: in the year 2000 this Sweet-clover was found to be widely distributed along the Alaska Highway between Johnson's Crossing and the Alaska border (13 specimens in DAO); gravel road- side, 14 km S of Jakes Corner, 60°19'05.4”"N 134°10'08.6"W, Cody & Cody 37388, 30 July 2000 (DAO); gravel clearing, Haines Highway Km 195.5, Dezadeash Lake Campground _ entrance, 60°23'52.9"N 137°02'51.3"W, Cody & Cody 37265, 28 July 2000 (DAO); dense stand along roadside, Faro, Douglass Drive off Lorna Blvd., 62°13'37.3"N 133°20'19.2"W, Cody & Cody 36981, 20 July 2000 (DAO); gravel at edge of road, Dempster Highway Km 1, 63°59'20.9"N 138°43'34"W, Cody & Cody 37018, 21 July 2000 (DAO); roadside gravel, Dempster Highway 4 km S of Benson Creek, 64°12'42.2"N 138°33'39.0"W, Cody & Cody 37024, 22 July 2000 (DAO). Cody et al. (1998, 2001) reported extensions of this widespread roadside species. The specimens cited above show a tremendous extension of this species in the southern half of the Territory in the year 2000. Melilotus officinalis (L.) Lam., Yellow Sweet-clover — YUKON: gravel clearing, Haines Highway Km 195.5, 60°23'50.9"N 137°02'51.3"W, Cody & Cody 37266, 28 July 2000 (DAO); roadside ditch, Robert Campbell Highway Km 477, 62°11'50.9"N 134°32'45.1"W, Cody & Cody 36985, 20 July 2000 (DAO); roadside gravel, Alaska Highway Km 1414.5, 60°28'18.1"N 134°15'43.6"W, Cody & Cody 37385, 30 July 2000 (DAO); roadside gravel, Alaska Highway, 60°48'38.2"N 135°13'31.2"W, Cody & 2002 Coby, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 467 Cody 37377, 29 July 2000 (DAO); disturbed gravel roadside, Alaska Highway, 77 km W of Klondike Highway junction, 60°47'30.4"N 136°29'45.2’W, Cody & Cody 37055, 24 July 2000 (DAO); gravel roadside, Alaska Highway Km 1590, 60°45’49.1"N 136°49'56.4"W, Cody & Cody 37373, 29 July 2000 (DAO); disturbed gravel by road, Alaska Highway, 1 km E of Haines Junction, 60°46'20.2"N 137°31'05.3"W, Cody & Cody 37061, 25 July 2000 (DAO); broad gravel roadside, Alaska Highway 400 m NW of White Creek bridge, 61°59'27.4"N 140°33'55.6"W, Cody & Cody 37150, 26 July 2000 (DAO); gravel beside highway, Alaska Highway 4.9 km SE of border crossing, 62°34'46.1"N 140°57'20.9"W, Cody & Cody 37220, 26 July 2000 (DAO). Although not nearly as common in the southern half of the Territory as M. alba, this yellow flowered species has also become widely distributed along the highways in the year 2000. Onobrynchis viciifolia Scop., Sainfoin — YUKON: height about 1 m in dense clump, roadside, Alaska Highway Km 1463, 7 km E of Robert Service Exit, Whitehorse, 60°37'59.4"N 134°59'52.3"W, Cody & Cody 37421, 31 July 2000 (DAO); Whitehorse, Kelley’s Canyon, Km 1440 Alaska Highway, 60°34.73'N 134°36.58'W, B. Bennett 00-876, 10 Sept. 2000 (DAO). In 1980 Cody and Ginns collected this introduced species adjacent to the Alaska Highway in the vicinity of Whitehorse. The specimens cited above confirm its occur- rence in the region. It was also noted eastward along the highway. Oxytropis campestris (L.) DC. ssp. varians (Rydb.) Cody, Field Locoweed — YUKON: common on river- bars, Wind River Camp #1, 64°40.39'N 134°35.96’W, B. Bennett 00-747, 2 July 2000 (B. Bennett Herbarium, photo DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 140 kilometers south of sites adjacent to the Peel River. Trifolium hybridum L., Alsike Clover — YUKON: roadside gravel, 24.5 road kilometers S of Carcross at Yukon/BC border, 60°00'00.6”N 134°39'47'W, Cody & Cody 37404, 30 July 2000 (DAO); roadside gravel, 17 km S of Jakes Corner, 64°19'42.1"N 134°11'45.9"W, Cody & Cody 37392, 30 July 2000 (DAO); disturbed gravel beside road, 103 km S of Carmacks on Klondike Highway, 61°15'33.4"N 135°28'45.5"W, Cody & Cody 37051, 24 July 2000 (DAO); gravel roadside, Top of the World Highway Km 2.6, 64°04'21.2”N 139°27'54.4"W, Cody & Cody 37029, 22 July 2000 (DAO); cleared gravel area, Top of the World Highway Km 56, 64°13'49.3"N 140°16'30.9"W, Cody & Cody 37042, 22 July 2000 (DAO); gravel roadside, Alaska Highway 21 km NW of Lake Creek Campsite, 61°58'33.8"N 140°27'50.3"W, Cody & Cody 37141, 26 July 2000 (DAO); broad gravel roadside, Alaska Highway 400 m NW of White River bridge, 61°59'27.4"N 140°33'55.6"W, Cody & Cody 37146 (DAO); disturbed gravel at road stop, Alaska High- way 7.4 km NW of Snag Junction campground, 62°17'16.4"N 140°46'45.5”"W, Cody & Cody 37178, 26 July 2000 (DAO); gravel roadside and ditch, Alaska Highway, 1.5 km N of Beaver Creek bridge, 62°22'36.2"N 140°52'35.6"W, 26 July 2000 (DAO); slope below rest area, Alaska Highway, 18 km SE of border crossing, 62°29'57.9"N 140°51.48.4”W, 26 July 2000 (DAO); gravel slope, Alaska Highway just east of border line, 62°36'52.8”N 140°59'59.8’"W, Cody & Cody 37202, 26 July 2000 (DAO). The specimens of this introduced species cited above extend the known distribution in the Territory (Cody 1996) to the Alaska border adjacent to the Alaska Highway, in the extreme south, and adjacent to the Top of the World Highway. Trifolium pratense L., Red Clover — YUKON: road- side gravel, Haines Highway Km 179, 60°15'55.4”N 136°58'30.4"W, Cody & Cody 37260, 28 July 2000 (DAO); disturbed gravel by roadside, Haines Highway Km 222, 100 mS of bridge over Kathleen River, 60°35’N 137°48.9"W, Cody & Cody 37270, 28 July 2000 (DAO). Cody (1996) knew this introduced species in the Territory from only seven sites. An additional site in the extreme southeast was reported by Cody et al. (2000). It is new to the Haines Highway although it was previously known from Haines Junction. Trifolium repens L., White Clover — YUKON: gravel roadside at rest stop, Haines Highway Km 150, 60°02'21.9"N 136°52'58.7"W, Cody & Cody 37252, 28 July 2000 (DAO); gravel by road stop, Haines Highway Km 173, 60°13'00.9"N 136°57'55.2’W, Cody & Cody 37257, 28 July 2000 (DAO). The specimens of this introduced species cited above are an extension of the known range in the Territory (Cody 1996) of about 125 kilometers southwest from the vicinity of Whitehorse. ELAEAGNACEAE Elaeagnus commutata Bernh., Silverberry — YUKON: cobbly silty riverbar with Salix alaxensis, rare, only place seen, Peel River between Snake and Bonnet Plume rivers, 65°58.61'N 134°48.61'W, B. Bennett 00-787, 10 July 2000 (DAO). This is the northernmost site yet found in the Territory (Cody 1996). It is an extension of about 115 kilometers northeast of sites adjacent to the Bonnet Plume River. Shepherdia canadensis (L.) Nutt., Soapberry — YUKON: open Picea glauca forest, Wind River Camp #1, 64°34.43'N 134°28.88'W, B. Bennett 00-227, 2 July 2000 (DAO); common on riverbar dominated by Salix alaxensis, Wind River Camp #4, 65°22.89'N 135°25.1'W, B. Bennett 00-257, 5 July 2000 (DAO); in young Populus balsamifera forest on riverbar, Peel River, Taco Bar, 66°00.15'N 134°13'W, B. Bennett 00-395, 13 July 2000 (DAO). The specimens cited above are extensions of the range in er 468 the Territory (Cody 1996) of about 175 kilometers to the south and 60 kilometers to the east of a site adjacent to lati- tude 66°N. APIACEAE (UMBELLIFERAE) Sium suave Walt., Water-parsnip — YUKON: lake- shore bordered by Picea glauca, Tatchun/ Frenchman Rd. Km 40, Tatchun Lake Campground, 62°16'50.0"N 136°02'09.3”W, Cody & Cody 37007, 21 July 2000 (DAO); wet shore of Gravel Lake, Klondike Highway Km 626, 63°48'26.5"N 137993" 56.7" W, Cady ‘& “Cody'37012 (DAO); Horseshoe Slough, 63°26’N 135°02'W, D. Mossop s.n., 13 July 2000 (B. Bennett Herbarium, photo DAO); Eagle Plains/Peel area, 66°24’N 136°58'W, L. Schroeder 17, 26 July 1994 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The nearest site to the first three specimens cited above is adjacent to the Nordenskiold River below Montague Mtn. (Cody et al. 2000). The fourth specimen cited above is the northernmost yet found in the Territory (Cody 1996). It is an extension north of the known range of about 300 kilometers from a site near the junction of the Klondike and Dempster highways. CORNACEAE Cornus canadensis L., Bunchberry — YUKON: ripar- ian Picea glauca/Populus balsamifera feathermoss forest, Peel River between Snake and Bonnet Plume rivers, 65°57.61'N 134°25.68”W, B. Bennett 00-703, 12 July 2000 (DAO); riparian Picea glauca/Populus balsamifera forest with thick moss cover, Peel River, 66°00.36'N 134°43.19'W, B. Bennett 00-463, 11 July 2000 (DAO). This species was not seen on the Wind River or farther upstream on the Peel River. The specimens cited above are the northernmost yet found in the Territory (Cody 1996). They are from sites about 300 kilometers north of a site just south of latitude 64°N. Cornus stolonifera Michx., Red-osier Dogwood — YUKON: Salix alaxensis/Alnus incana/Rosa acicu- laris/Shepherdia canadensis/Dryas drummondiana gravelly riverbar, Wind River Camp #6, 65°40.46’N 135°11.76'W, B. Bennett 00-848, 7 July 2000 (DAO); along riverbars under Salix alaxensis and Alnus incana thickets, confluence of Wind and Peel rivers, 65°50.48'N 135°18.25'W, B. Bennett 00-111, 9 July 2000 (DAO); moist Picea glauca/Alnus thick- et, riparian seeps, Peel River Site #1, 65°58.06’N 134°49.69'W, B. Bennett 00-432, 10 July 2000 (DAO); in young Populus balsamifera forest on rivetbar, Peel ‘River, Taco Bar, 60°00:15’N 134°13'W, B. Bennett 00-379, 13 July 2000 (DAO). The specimens cited above are near sites adjacent to the lower Bonnet Plume River (Cody et al. 2000) including the northernmost yet found in the Territory (Cody 1996) col- lected about 50 kilometers to the northeast. PYROLACEAE Moneses uniflora (L.) Gray, One-flowered Pyrola — YUKON: older Picea glauca/Populus balsamifera for- est beside creek in shady mossy area, Wind River Site THE CANADIAN FIELD-NATURALIST _—_— ara 0 Tee a Vol. 116 #1, 64°40.39'N 134°35.96'W, B. Bennett 00-743, 2 July 2000 (DAO); growing in moss on river bank, Snake River, 64°47'N 132°40'W, J. Meikle 99-037, 18 July 1999 (Yukon Renewable Resources, photo DAO); steep slope with Populus tremuloides, Wind River, Deception Mtn., 65°36.03'N 135°28.51'W, B. Bennett 00-443, 7 July 2000 (DAO); riparian Picea glauca/Populus balsamifera feathermoss forest, Peel River between Snake and Bonnet Plume rivers, 65°57.68'N 134°25.68’W, B. Bennett 00-702, 12 July 2000 (DAO). The specimens cited above from between latitude 64°and 66°N are intermediate between sites mapped by Cody (1996) adjacent to the Dempster Highway and the upper Bonnet Plume River. Orthilia secunda (L.) House, One-sided Wintergreen — YUKON: in young Populus balsamifera forest on riverbar, Peel River, Taco Bar, 66°00.15'N 134°13'W, B. Bennett 00-394, 13 July 2000 (DAO); riparian Picea glauca/Populus balsamifera/Alnus incana forest, Wind River, Deception Mountain, 65°36.03'N 135°28.41'W, B. Bennett 00-416, 7 July 2000 (DAO); tussock tundra, Wind River Camp #2, 64°51.83'N 134°38.85'W, B. Bennett 00-829, 4 July 2000 (DAO); older Picea glauca/Populus balsam- ifera forest beside creek in shady mossy area, Wind River, 64°40.39’N 134°35.96'W, B. Bennett 00-745, 2 July 2000 (DAO). The specimens cited above are from between sites north of 66°N and the upper part of the Bonnet Plume River (Cody 1996). Pyrola asarifolia Michx., Pink-flowered Winter- green — YUKON: older Picea glauca/Populus bal- samifera forest beside creek in mossy shady area, 64°40.39'N 134°35.96'W, B. Bennett 00-720, 2 July 2000 (DAO); riverbar, Wind River Camp #6, 65°40.46'N 135°11.76'W, B. Bennett 00-146, 7 July 2000 (DAO); moist Picea glauca/Alnus thicket ripar- ian with seeps, Peel River Site #1, 65°58.06'N 134°49.69'W, B. Bennett 00-401, 10 July 2000 (DAO); in young Populus balsamifera riparian forest on riverbar, Peel River, Taco Bar, 66°00.15’N 134°13'W, B. Bennett 00-392, 13 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 175 kilometers east of the Dempster Highway between latitudes 64°N and 67°N. Pyrola chlorantha Sw., Greenish Wintergreen — YUKON: riparian Picea glauca/Populus balsamiferal Alnus incana forest, Wind River, Deception Mountain, 65°36.03'N 135°28.41'W, B. Bennett 00- 421, 7 July 2000 (B. Bennett Herbarium, photo DAO); riparian Picea glauca/Populus balsamifera forest with thick moss cover, Peel River, 66°00.36'N 134°43.19'W, B. Bennett 00-468, 11 July 2000 (DAO); wet moss under spruce, Alaska Highway, 2.3 km SE of border crossing, 62°36'05.2”N 140°58'04.9"W, Cody & Cody 37215, 26 July 2000 (DAO). 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 469 The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 75 kilo- meters east of sites just north of latitude 66°N and of about 200 kilometers northwest of a site near the south end of Kluane Lake. Pyrola grandiflora Radius, Arctic Wintergreen — Yukon: steep northwest-facing slope of lower cirque basin, Bonnet Plume River, 64°51'19"N 133°27'53”W, V. Loewen 99-6-26, 11 July 1999 (DAO); in moist White Spruce spruce forest with moss understory, Snake River, J. Meikle 99-038, 19 July 1999 (Yukon Renewable Resources, photo DAO). | The specimens cited above extend the known range in the Territory (Cody 1996) about 75 kilometers north of a site in the upper Bonnet Plume River. ERICACEAE Arctostaphylos rubra (Rehd. & Wils.) Fern., Red Alpine Bearberry — YUKON: open gravelly riverbar, Wind River, 64°40.39'N 134°35.96'W, B. Bennett 00-180, 3 July 2000 (DAO); hummocky Picea mari- ana site south of recent fire west of cat trail, Wind River, 64°58.67'N 134°47.12’W, B. Bennett 00-298, 4 July 2000 (DAO); moderately steep east-facing subalpine slope, Bonnet Plume River east of Fairchild Lake, 64°58'03”N 133°41'12’W, V. Loewen 99-4-14, 11 July 1999 (DAO); convex mid- slope, rock/ soil/Dryas ridge, Upper Bonnet Plume River Drainage Site #128, J. Staniforth 00-107, 8 July 2000 (DAO). The specimens cited above extend the known range in the Territory (Cody 1996) about 200 kilometers east of sites adjacent to the Hart River. Arctostaphylos uva-ursi (L.) Spreng., Common Bearberry — YUKON: on steep vegetated slope above river, Wind River, 64°40.39'’N 134°35.96'’W, B. Bennett 00-737, 2 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 150 kilometers east of a site between the Blackstone and Hart rivers. Chamaedaphne calyculata (L.) Moench., Leatherleaf — YUKON: riverbar, Peel River, 65°56.03'N 134°58.84'’W, B. Bennett 00-780, 9 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 50 kilometers northeast of a site between the Hart and Wind rivers. Ledum groenlandicum Oeder, Labrador-tea — YUKON: in hummocky Picea/Salix forest near river, Wind River, McClusky Lake, 64°34.19'N 134°25.77'W, B. Bennett 00-377, 2 July 2000 (DAO). The specimen cited above is intermediate between sites mapped by Cody (1996) adjacent to the Hart River and the upper Bonnet Plume River. Oxycoccus microcarpus Turcz., Bog Cranberry — YUKON: boggy area with open pools, Larix laricina/ Rubus chamaemorus/Ledum groenlandicum/Betula papyrifera, Peel River, 66°00.36’N 134°43.19’'W, B. Bennett 00-465, 11 July 2000 (DAO); fen ingrowing with Carex limosa and Sphagnum with Andromeda, Vuntut National Park, B. Bennett 00-560, 1 Aug. 2000 (DAO). The specimen first cited above is an extension of the known range in the Territory of about 55 kilometers east of sites mapped by Cody (1996) from just north of latitude 66°N; the second specimen, from north of 68°N, is notable since there are only two other sites north of this latitude. PRIMULACEAE Androsace chamaejasme Host. ssp. lehmanniana (Spreng.) Hultén, Rock-jasmine — YUKON: south- west-facing slope, small carbonate face, Snake River, 65°06.5’N 133°07'W, J. Meikle 99-041A, 19 July 1999 (Yukon Renewable Resources, photo DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 50 kilometers east of a site adjacent to the Bonnet Plume River. Primula mistassinica Michx., Bird's-eye Primrose — YUKON: riverbar, Wind River, 64°34.43'N 134°28.88'W, B. Bennett 00-211, 2 July 2000 (DAO); riverbar, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-356, 2 July 2000, (DAO); in moss on riverbank, Wind River Camp #5, 65°35.38'N 135°27.6'W, B. Bennett 00-141, 6 July 2000 (DAO); common on turfy sites by creek in woods below timberline, Upper Big Creek, Dawson Range, 62°18'N 137°53’W, R. Frisch s.n., 7 June 1982 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The first three specimens cited above are the northernmost yet known in the Territory (Cody 1996). They are an extension of the known range of about 225 kilometers northwest from a site adjacent to the upper Bonnet Plume River. The fourth specimen is an extension of the known range of about 80 kilometers east of a site west of longitude 139°W. PLUMBAGINACEAE Armeria maritima (Mill.) Willd. ssp. arctica (Cham.) Hultén, Sea Thrift — YUKON: boggy tussock tundra near small pond, uncommon in this area but several groups seen throughout and onto riverbars, Wind River Camp #2, 64°51.83'’N 134°38.85'W, B. Bennett 00-818, 4 July 2000 (DAO); base of talus slope in calcareous outwash, Wind River Camp #3 east bank near confluence of Royal Creek, 65°06.83'N 134°47.12'W, B. Bennett 00-216, 4 July 2000 (B. Bennett Herbarium, photo DAQ). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 575 kilometers south of the Arctic coast and British Mountains. It is how- ever known in the Mackenzie Mountains to the east. MENYANTHACEAE Menyanthes trifoliata L., Buckbean — YUKON: Sam Lake, Vuntut National Park, 68°24.24'N 138°37.58'W, B. Bennett 00-552, 5 Aug. 2000 (DAO). _ ) ans: i SF ih athe 1 470 The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 50 kilometers northeast of the only other collection from north of latitude 68°N. POLEMONIACEAE Phlox alaskensis Jordal, Alaskan Phlox — YUKON: in steep loose talus, west-facing slope, Wind River Camp #3, 65°06.83’N 134°37.12'W, B. Bennett 00- 157, 00-217, 4 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 80 kilo- meters east of a site east of Hart River. Polemonium boreale Adams, Northern Jacob’s-lad- der — YUKON: gravel beside Alaska Highway, Beaver Creek Community, 62°23'11.1”N 140°52'28.3"W, Cody & Cody 37227 (DAO). The nearest site of this species adjacent to the Alaska Highway to the specimen cited above is about 140 kilome- ters to the southeast near the north end of Kluane Lake (Cody 1996). It is however known from adjacent to the Alaska border at about latitude 62°N. BORAGINACEAE Mertensia paniculata (Ait.) G. Don var. paniculata, Bluebell — YUKON: borders of roads in campground by Salix, Alnus and Picea, Pine Valley Campground, Alaska Highway, 61°51'14.3"N 140°08'58.9"W, Cody & Cody 37114, 26 July 2000 (DAO); gravel beside Alaska Highway, Beaver Creek Community, 62°23'11.1"N 140°52'28.3"W, Cody & Cody 37226 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 180 kilometers northwest of a site adjacent to the south end of Kluane Lake. LAMIACEAE (LABIATAE) Scutellaria galericulata L. var. pubescens Benth., Narrow-leaved Skullcap — YUKON: wet shore of Gravel Lake, Klondike Highway Km 626, 63°48'26.5"N 137°53'56.7"W, Cody & Cody 37010, 21 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory of about 100 kilometers to the west from the vicinity of Mayo (Cody 1996). Douglas et al. (1981) considered this species rare in the Territory. SCROPHULARIACEAE Pedicularis capitata Adams, Capitate Lousewort — YUKON: lower South Bend on Snake River, 65°06’N 133°06'W, J. Meikle 99-045, 19 July 1999 (Yukon Renewable Resources, photo DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 50 kilometers northeast of a site between the Wind and Bonnet Plume rivers. Pedicularis langsdorfii Fisch. ssp. arctica (R.Br.) Pennell ex Hultén, Langdorf’s Lousewort — YUKON: riverbar, shrubby Salix, occasional, Wind River Outfitter’s Camp #1, 64°34.43'N 134°28.88'W, B. Bennett 00-266, 2 July 2000 (DAO); open dolomite THE CANADIAN FIELD-NATURALIST Vol. 116 delta, Wind River, 64°48.46’N 134°41.34’W, B. Bennett 00-319, 3 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 80 kilo- meters north of a site just north of latitude 64°N. Pedicularis verticillata L., Whorled Lousewort — YUKON: riverbar, Wind River, 64°34.43’N 134°28.88'W, B. Bennett 00-269, 2 July 2000 (DAO); common on riverbar dominated by Salix alaxensis, Wind River Camp #1, 64°40.39'N 134°35.96’W, B. Bennett 00-249, 3 July 2000 (DAO). — The specimens cited above are intermediate between sites west of the Hart River mapped by Cody (1996) and a site adjacent to the Snake River (Cody et al. 2001). LENTIBULARIACEAE Utricularia vulgaris L. ssp. macrorhiza (Le Conte) Clausen, Greater Bladderwort — YUKON: in shal- lows of small upland lake, Frances Lake, 61°11'N 129°08'W, R. Rosie 2007, 5 June 1998 (DAO), R. Rosie 2008, 17 Oct. 1999 (DAO). The specimens cited above are intermediate between sites adjacent to the Canol Road (Cody 1996) and a site adjacent to the Beaver River (Cody et al. 2000). PLANTAGINACEAE Plantago eriopoda Torr., Saline Plantain — YUKON: rare along roadside, Klondike Highway 81 km S of Stewart Crossing, 62°44'52.6"N 136°39'32.4"W, Cody & Cody 37046, 24 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 100 kilometers north of sites adjacent to the highway south of Carmacks. Plantago major L., Common Plantain — YUKON: dis- turbed gravel by boat ramp, North Canol Road Km 333 at Dragon Lake, 62°33'19.6’N 131°20'26.8"W, Cody & Cody 36947, 19 July 2000 (DAO); gravel cleared area, Faro, 62°13'59.2"N 133°20) 373 ee Cody & Cody 36967, 20 July 2000 (DAO); gravel by roadside stop, Little Salmon River, Campbell Highway Km 553, 62°05’N 135°30'W, Cody & Cody 35807, 5 July 1999 (DAO); Koidern River (Edith Creek) behind the Pine Valley Campground, 61°51'19.6"N 140°09'11.2"W, Cody & Cody 37123, 26 July 2000 (DAO); gravel slope, Alaska Highway, just east of Alaska border line, 62°36'52.8"N 140°59'59.8"W, Cody & Cody 37201, 26 July 2000 (DAO); gravel parking area beside shrubs at Forty Mile Herd Rest Stop, Top of the World Highway Km 14, 64°05'16.5"N 139°36'49.4"W, Cody & Cody 37032, 22 July 2000 (DAO). This introduced species which is widespread in the Territory (Cody 1996) was not previously known from adjacent to the Canol Road, Campbell Highway, extreme northern Alaska Highway or Top of the World Highway. CAPRIFOLIACEAE _Linnaea borealis L. ssp. americana (Forbes) Hultén, Twinflower — on steep vegetated slope above river, Wind River, 64°40.39'N 134°35.96’W, B. Bennett 00- 725, 2 July 2000 (DAO); riparian Picea 2002 Copy, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 471 glauca/Populus balsamifera/Alnus incana forest, Wind River, Deception Mountain, 65°36.03'N 135°28.41'W, B. Bennett 00-281, 7 July 2000 (DAO); riparian Picea glauca, Populus balsamifera forest with thick moss cover, Peel River, 66°00.36'N 133°43.19'W, B. Bennett 00-466, 11 July 2000 (DAO). The specimens cited above extend the known distribu- tion in the Territory (Cody 1996) about 150 kilometers to the south and 75 kilometers east of a site between the Hart and Wind rivers. Viburnum edule (Michx.) Raf., Low Bush-cranberry — YUKON: lush Picea glauca/Alnus incana estab- lished riparian thicket, Wind River Camp #4, 65°22.89'N 135°26.1'W, B. Bennett 00-168, 5 July 2000 (DAO); riparian Picea glauca, Populus bal- samifera, Alnus incana forest, Deception Mountain, Wind River, 65°36.03’N 135°28.41'W, B. Bennett 00-428, 7 July 2000 (DAO); edge of river in older Salix alaxensis/S. pseudomonticola/S. pulchral/ Calamagrostis canadensis meadow, 65°58.13'N 134°25.68'W, B. Bennett 00-714, 12 July 2000 (DAO); older riparian Picea glauca/Populus balsam- ifera forest with Eleaganus, Peel River Camp #9, 65°58.13'N 134°48.61'W, B. Bennett 00-792, 10 July 2000 (DAO); shrub-thicket on strongly mound- ed lower slope, Upper Bonnet Plume River Drainage near site #151, 64°30'24"N 133°01'59"W, J. Staniforth 00-144, 11 July 2000 (DAO). The specimens cited above are an extension of the range in the Territory (Cody 1996) of about 250 kilometers north and 100 kilometers northeast from sites south of latitude 64°N. CAMPANULACEAE Campanula aurita Greene, Yukon Bellflower — in calcareous talus, Peel River Site #1, 65°58.06'N 134°49.69'W, B. Bennett 00-407, 10 July 2000 (B. Bennett Herbarium, photo DAO); riverbar, base of cliff, Wind River, 65°12.49’N 135°13.17'W, B. Bennett 00-365, 5 July 2000 (B. Bennett Herbarium, photo DAO); calcareous scree, Wind River, 64°51.83’N 134°38.85'W, B. Bennett 00-822, 4 July 2000 (DAO); open dolomite area, Wind River, 64°48.46'N 134°41.46'W, B. Bennett 00-317, 3 July 2000 (DAO). The specimens cited above extend the known range in the Territory (Cody 1996) about 165 kilometers to the south from a site north of latitude 66°N. Campanula uniflora L., Arctic Harebell — YUKON: moist seepage area, Bonnet Plume River, 64°53'27"N 133°48'01"W, V. Loewen 99-19-73, 13 July 1999 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 185 kilometers east of a site west of Hart River. ASTERACEAE (COMPOSITAE) Achillea millefolium L. ssp. lanulosa (Nutt.) Piper, Common Yarrow — YUKON: roadside gravel, Burwash Landing, Alaska Highway, 61°21'17.5’N 139°00'04.6"W, Cody & Cody 37094, 25 July 2000 (DAO); borders of roads in campground by Salix, Alnus and Picea, Pine Valley Campground, Alaska Highway, 61°51'14.3"N 140°08'58.9"W, Cody & Cody 37107, 26 July 2000 (DAO); open riverbar, recently flooded, Peel River, 65°55.97'N 135°03'W, B. Bennett 00-103, 9 July 2000 (B. Bennett Her- barium, photo DAO). Douglas et al. (1981) considered this species rare in the Territory. Cody (1996) knew it from nine sites north to lati- tude 64°N. The first specimens cited above extend the known range in the Territory about 290 kilometers north- west of Whitehorse. The last specimen that is the northern- most yet found in the Territory is an extension of the known range to the north of about 230 kilometers from a site in the vicinity of Mayo. Achillea sibirica Ledeb., Siberian Yarrow — YUKON: gravel slope, Alaska Highway just east of Alaska bor- der line, 62°36'53.8”N 140°59'59.8"W, Cody & Cody 37198, 26 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 100 kilometers to the northwest from a site south of latitude 62°N adjacent to the Alaska Highway. Antennaria monocephala DC. ssp. monocephala, Pygmy Pussytoes — YUKON: saddle in alpine upper slope of mountain east of Fairchild Lake, Bonnet Plume River, 64°58'23"N 133°42'15”W, V. Loewen 99-1-4, 11 July 1999 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 80 kilometers east of a site west of longitude 135°W. Antennaria pulcherrima (Hook.) Greene, Showy Pussytoes — YUKON: common on riverbar, Wind River Camp #1, 64°40.39'N 134°35.96'W, B. Bennett 00-247, 3 July 2000 (B. Bennett Herbarium, photo DAO). The specimen cited above is about 100 kilometers north- west of a site adjacent to the upper Bonnet Plume River (Cody 1996) and 100 kilometers southwest of the northern- most site in the Territory (Cody et al. 2001). Artemisia alaskana Rydb., Alaska Sagebrush — YUKON: cobbly silty riverbar with Salix alaxensis shrub, Peel River Camp #9 between Snake and Bonnet Plume rivers, 65°58.13'N 134°48.61'W, B. Bennett 00-788, 10 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 125 kilometers north of a site adjacent to the Bonnet Plume River. Artemisia biennis Willd., Wormwood — YUKON: gravel cleared area, Faro, 62°13'59.2"N 133°20'57.3"W, Cody & Cody 36970, 20 July 2000 (DAO). The specimen of this introduced species cited above is only from the third known area in the Territory. It was known to Cody (1996) only from the vicinity of the Porcupine River and Cody et al. (2000) from the extreme southeast. ee 472 Artemisia campestris L. s.l., Northern Wormwood — YUKON: riverbar, Wind River, 64°44.64'N 134°38.3'W, B. Bennett 00-181, 3 July 2000 (DAO); gravelly riverbar, Wind River, 65°35.38'N 135°27.6'W, B. Bennett 00-140, 6 July 2000 (DAO). The specimens cited above are intermediate between the two northernmost sites mapped by Cody (1996) adjacent to the Bonnet Plume River and north of latitude 66°N. Artemisia michauxiana Bess. in Hook., Michaux’'s Mugwort — YUKON: only plant growing in railway bed, Quartz Road, Whitehorse, 60°44.05'N 135°04.09’W, B. Bennett 00-052, 6 June 2000 (B. Bennett Herbarium, photo DAO); very steep loose shale, Tagish Road hill across from Crag Lake 60°15'24”N 134°29'12”W, B. Bennett 01-016, 19 May 2001 (B. Bennett Herbarium, photo DAO). This species was considered rare in the Territory (Douglas et al. 1981) on the basis of a collection from the vicinity of Johnsons Crossing; Cody et al. (2001) reported a second collection from a sand beach at Carcross; the first specimen cited above is the northernmost yet found in the Territory; the second specimen cited above, which is only from the fourth site known in the .Territory is from a site northeast of Carcross. Artemisia tilesii Ledeb. s.1., Aleutian Mugwort — YUKON: in talus below cliff, Wind River, 65°12.49’N 135°13.17'W, B. Bennett 00-135, 5 July 2000 (DAO). The specimen cited above is between sites mapped by Cody (1996) from the Hart River and Dempster Highway and a site adjacent to the Snake River (Cody et al. 2001). Aster sibiricus L., Arctic Aster — YUKON: semi- active flood plain on gravelly soil in back-channel, Snake River, 64°45.57’N 132°35.81'W, J. Meikle 99-051, 18 July 1999 (Yukon Renewable Resources, photo DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 75 kilometers northwest of a site adjacent to the upper Bonnet Plume River. Crepis nana Richards., Dwarf Hawksbeard — YUKON: roadside gravel slope, Alaska Highway, 20 km NW of White River bridge, 62°08'39.6”N 140°41'05.0"W, Cody & Cody 36157, 26 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory of about 120 kilometers from a site near the northern end of Kluane Lake (Cody 1996). Erigeron hyssopifolius Michx., Hyssop-leaf Flea- bane — YUKON: common on riverbars, Wind River Camp #1, 64°40.39'N 134°35.96’W, B. Bennett 00- 353, 00-756, 2 July 2000 (DAO); riverbar, Wind River Camp #4, 65°22.89'N 135°26.1’W, B. Bennett 00-301, 5 July 2000 (DAO); vegetated established river bank, Wind River Camp #6, 65°40.46’N 135°11.76'W, B. Bennett 00-142, 7 July 2000 (DAO); silty seep on edge of river, Peel River Site #1, 65°58.06'N 134°49.69'W, B. Bennett 00-434, 10 July 2000 (DAO). THE CANADIAN FIELD-NATURALIST ae | Vol. 116 - Douglas et al. (1981) considered this species rare in the Territory in the Richardson Mts. but the approximate circle on his map was too far south. Cody (1996) knew it from four sites between latitudes 64°N and 66°N from adjacent to the Bonnet Plume River and between the Wind and Hart rivers. The specimens cited above are the first known from the Wind and Peel rivers. Erigeron lonchophyllus Hook., Spear-leaved Flea- bane — YUKON: gravelly roadside, Tatchun/ Frenchman Road, 62°07'N 135°45’W, B. Bennett 99-127, 15 July 1999 (DAO). The specimen cited above is intermediate between sites mapped by Cody (1996) just north of Whitehorse and in the vicinity of Mayo. Haplopappus macleanii Brandegee, Northern Mock Goldenweed — YUKON: dry open steep hillside just east of pullout on Robert Campbell Highway, Carmacks, Eagle Rock, 62°01.75'’N 135°50.62’W, B. Bennett 00-048, 20 May 2000 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is from a site intermediate between the north end of Lake Laberge and the Klondike Highway south of Pelly Crossing (Cody 1996). Matricaria matricarioides (Less.) Porter, Pineapple- weed — YUKON: gravel roadside, Koidern River (Edith Creek) behind the Pine Valley Campground, 61°51'19.6"N 140°09'11.2"W, Cody & Cody 37130, 26 July 2000 (DAO); disturbed gravel at roadstop, Alaska Highway 7.4 km NW of Snag Junction Campground, 62°16'16.4"N 140°46'45.5"W, Cody & Cody 37187A, 26 July 2000 (DAO); disturbed camp area, Wind River Camp #2, 64°45.05’'N 134°38.75'W, B. Bennett 00-186, 3 July 2000 (DAO). The specimens cited above extend the known distribu- tion of this cosmopolitan weed, which is widespread in the Territory (Cody 1996), about 75 kilometers to the north- west adjacent to the Alaska Highway near the Alaska bor- der and to the northeast about 115 kilometers from the vicinity of Mayo. Matricaria perforata Mérat, Scentless Mayweed — YUKON: sandy gravel cleared area, Top of the World Highway Km 32, 64°10'26.6"N 139°52’38.3"W, Cody & Cody 37040, 22 July 2000 (DAO). This adventive species from Europe was only known in the Territory (Cody 1996) from the vicinities of Ross River and Frances Lake and to Cody et al. (2000) from the vicini- ty of Colwell Bay. Petasites frigidus (L.) Fries ssp. frigidus, Sweet Coltsfoot — YUKON: moist area between cliff and Salix/Populus/Picea forest, Wind River Camp #4, 65°22.89'N 135°26.1'W, B. Bennett 00-200, 5 July 2000 (DAO); steep slope with Populus tremuloides, Wind River, Deception Mtn., 65°36.03'N -135°28.41'W, B. Bennett 00-466, 7 July 2000 (DAO); McClusky Lake, Wind River, 64°54.19'N 134°25.77'W, B. Bennett 00-326, 2 July 2000 (DAO). 2002 Coby, KENNEDY, BENNETT, AND LOEWEN: VASCULAR PLANTS IN YUKON IV 473 The specimens cited above are from sites intermediate between a site adjacent to the Blackstone River (Cody 1996) and the Snake River (Cody et al. 2001). Petasites frigidus (L.) Fries ssp. palmatus (Ait.) Cody — YUKON: slumping eroding banks, cobbles and silty muck, Wind River, 65°47.45’N 135°13'W, B. Bennett 00-121, 8 July 2000 (DAO); lush seep draining upland, Wind River, 65°14.8'N 135°18.51’W, B. Bennett 00-130, 5 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 500 kilometers from sites south of 61°30’N. Senecio atropurpureus (Ledeb.) Fedtsch. ssp. frigidus (Richards.) Hultén, Purple-haired Groundsel — YUKON: moderately steep north-facing slope, Bonnet Plume River, 64°51'10"”N 133°27'41”"W, V. Loewen 99-5-25, 11 July 1999 (DAO); steep talus slope, Wind River Camp #4, 65°22.89'N 135°26.1"W, B. Bennett 00-202, 5 July 2000 (B. Bennett Herbarium, photo DAO); Larix laricina/ Picea mariana/Sphagnum bog, Peel River Camp #10, 65°57'N 134°25.68'W, B. Bennett 00-708, 12 July 2000 (B. Bennett Herbarium, photo DAO); undulating spruce/shrub/low shrub/forb/ moss, Upper Bonnet Plume River Drainage Site #116, 64°26'6"N 132°15'28"W, J. Staniforth 00-015, 5 July 2000 (DAO); undulating upper slope, steep alpine ridge, rock rubble, bare soil and sparse vegetation, Upper Bonnet Plume River Drainage Site #125, 64°32'45"N 132°50'48"W, J. Staniforth 00-102, 8 July 2000 (DAO); Snake River, 65°01'N 133°07'W, J. Meikle 99-056, 19 July 1999 (Yukon Renewable Resources, photo DAO). The specimens cited above are intermediate between lat- itudes 64° and 66° east of a site just east of longitude 137°W (Cody 1996). Solidago canadensis L. var. salebrosa (Piper) Jones, Canada Goldenrod — YUKON: loose talus at base of steep shale scree slope, confluence of Peel and Wind rivers, 65°50.48'N 135°18.25'W, B. Bennett 00-784, 8 July 2000 (B. Bennett Herbarium, photo DAO); riverbar near slough, Peel River Camp #8, 65°56.03'N 134°58.43'W, B. Bennett 00-777, 9 July 2000 (DAO); cobbly silty riverbar with Salix alaxen- sis shrub, between Snake and Bonnet Plume rivers, 65°58.13'N 134°48.61'W, B. Bennett 00-794, 10 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 280 kilometers to the northwest from sites adjacent to the Yukon River in the vicinity of Dawson City. Solidago simplex Kunth, Spike-like Goldenrod — YUKON: less active areas on riverbar, Wind River, 65°12.49'N 135°13.17'W, B. Bennett 00-456, 5 July 2000 (B. Bennett Herbarium, photo DAO); slumping eroding banks, cobbles and silty muck, Wind River, 65°47.45'N 135°13'W, B. Bennett 00-123, 8 July 2000 (DAO); loose talus at base of steep active shale scree slope, junction of Wind and Peel rivers, 65°50.48'N 135°18.25'W, B. Bennett 00-783, 8 July 2000 (DAO). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 200 kilometers northeast from a site adjacent to the upper Bonnet Plume River. Taraxacum ceratophyllum (Ledeb.) DC., Horned Dandelion — YUKON: in open meadow near outfitter cabins, Wind River, McClusky Lake, 64°34.19’N 134°25.77"W, B. Bennett 00-340, 2 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 100 kilometers northwest of a site adjacent to the upper Bonnet Plume River. Taraxacum officinale Weber ex Wiggers, Common Dandelion — YUKON: open meadow near outfitter’s camp, Wind River, McClusky Lake, 64°34.19'N 134°25.77'W, B. Bennett 00-235, 2 July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 90 kilometers northeast from a site in the vicinity of Mayo. Acknowledgements We thank Gerald A. Mulligan for the identification of Brassicaceae (Cruciferae) specimens; George Argus for the identification of Salix specimens; Stephen J. Darbyshire for the identification of Elymus trachycaulus ssp. novae-angliae; John Meikle, Dave Mossop, Rhonda Rosie and Stu Whithers for their contributions; David Cody for assisting the senior author with collections in the summer of 2000 in the southern Yukon; artists Valerie Fulford and Lee Mennell; Paul Catling for reviewing an earlier version of this manuscript, and especially Leslie Durocher, for the many hours inputting this information on her computer. Literature Cited Black, M. L. 1944. Yukon Wild Flowers. Price, Temple- ton Syndicate, Vancouver, British Columbia. 96 pages. Brunton, D. F., and D. M. Britton. 1999. Maritime Quillwort, /soetes maritima (Isoetaceae), in the Yukon Territory. Canadian Field-Naturalist 113: 641-645. Cody, W. J. 1954. A History of Tillaea aquatica (Crassu- laceae) in Canada and Alaska. Rhodora 56: 96-101. Cody, W. J. 1994. The Flora of the Yukon Territory: Additions, Range Extensions and Comments. Canadian Field-Naturalist 108: 428-476. Cody, W. J. 1996. Flora of the Yukon Territory. National Research Council (NRC) Press, Ottawa, Ontario, Canada. 643 pages. Cody, W. J. 2000. Flora of the Yukon Territory, Second Edition. National Research Council (NRC) Press, Ottawa, Ontario, Canada. 669 pages. Cody, W. J., C. E. Kennedy, and B. Bennett. 1998. New Records of Vascular Plants in the Yukon Territory. Canadian Field-Naturalist 112: 289-328. 474 Cody, W. J., C. E. Kennedy, and B. Bennett. 2000. New Records of Vascular Plants in the Yukon Territory, II. Canadian Field-Naturalist 114: 417-443. Cody, W. J., C. E. Kennedy, and B. Bennett. 2001. New Records of Vascular Plants in the Yukon Territory, LI. Canadian Field-Naturalist 115: 301-322. Douglas, G. W., G. W. Argus, H. L. Dickson, and D. F. Brunton. 1981. The rare vascular plants of the Yukon. Syllogeus 28: 1-96. Douglas, G. W., G. B. Straley, and D. Meidinger. 1989. The vascular plants of British Columbia: Part 1 — Gymnosperms and dicotyledons (Aceraceae through Cucurbitaceae). British Columbia Ministry of Forests, Special Report Series 1. 208 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1990. The vascular plants of British Columbia: Part 2 — Dicotyledons (Diapenseaceae through Portulacaceae). British Columbia Ministry of Forests, Special Report Series 2. 158 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1991. The vascular plants of British Columbia: Part 3 — Dicotyledons (Primulaceae through Zygophyllaceae). British Columbia Ministry of Forests, Special Report Series 3. 177 pages. THE CANADIAN FIELD-NATURALIST = | Vol. 116 Douglas, G. W., G. B. Straley, and D. Meidinger. 1994. The vascular plants of British Columbia: Part 4 — Monocotyledons. British Columbia Ministry of Forests, Special Report Series 4. 257 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998. Illustrated Flora of British Columbia, Volume 2. British Columbia Ministry of Forests, Victoria, British Columbia, Canada. 401 pages. Hultén, E. 1950. Flora of Alaska and Yukon. Lunds Universitets Arsskrift. N.F. Avd. 2 Bd 46. Nr 1. Hultén, E. 1968. Flora of Alaska and Neighboring Terri- tories. Stanford University Press, Stanford, California. 1008 pages. Porsild, A. E. 1951. Botany of southeastern Yukon adja- cent to the Canol Road. National Museum of Canada Bulletin 121. 400 pages. Rosie, R. 1991. Range extensions and rare vascular plants from southeastern Yukon Territory. Canadian Field- Naturalist 105: 314-324. Sheviak, C. J. 1993. Cypripedium parviflorum Salib. var. makasin (Farwell) Sheviak. American Orchid Society Bulletin 64(4): 403. Received 11 November 2001 Accepted 23 July 2002 Breeding Birds in Forestry Plantations and Natural Forest in the Vicinity of Fundy National Park, New Brunswick Grec A. M. JoHNSoN! and BILL FREEDMAN2 'Jacques Whitford Environment Limited, P.O. Box 1116, Fredericton, New Brunswick E3B 5C2 Canada *Department of Biology, Dalhousie University, Halifax, Nova Scotia, B3H 4J1 Canada [Author to whom correspondence should be addressed] Johnson, Greg A. M., and Bill Freedman. 2002. Breeding birds in forestry plantations and natural forest in the vicinity of Fundy National Park, New Brunswick. Canadian Field-Naturalist 116(3): 475-487. We studied the populations of breeding birds in five reference stands of natural mixedwood forest and 11 conifer plantations up to 21-years old in southern New Brunswick, Canada. Variations of the distribution and abundance of birds were related to changes in the plant-species composition and structural attributes of their habitat. Bird species occurred in plantations in lev- els of abundance and diversity similar to that of reference forest, although community composition was highly dissimilar. Species of reference stands were typical of mature, mixedwood forest. Species of younger plantations were typical of open, early successional, upland habitats. Once the plantations became older than 13 years and the tree height exceeded about 5 m, birds typical of conifer forest began to invade the habitat, resulting in a mixed-species composition. Although cavity-con- taining snags were rare in the plantations, if they did occur in them or near their edge they were used by hole-nesting birds. Although we could not study a complete rotation, the field data and habitat trajectory suggest that the plantations will not support some elements of the avian biodiversity of the natural forest in the study area. The mature plantations will be highly depauperate in coarse-woody debris, snags, and cavities, and will not support species dependent on these critical habitat ele- ments. In addition, birds requiring habitat containing trees of larger size and of various species, including hardwoods, will not find the mature plantations to be suitable. To accommodate the needs of species potentially at risk from the extensive development of plantations, the following changes in forestry management practices should be instituted: (1) retention of natural cavity-trees, snags, coarse-woody debris, and hardwood trees and shrubs in residual non-harvested “islands” within clear-cuts and plantations; (b) retention of a patchy angiosperm component by leaving some areas untreated during herbicide applications; and (c) setting aside large areas of natural forest as non-harvested protected areas. Although we believe that these mitigations would help sustain elements of indigenous biodiversity that are at risk in an extensive industrial forest, we stress that these predictions would have to be tested through additional research and monitoring. Key Words: Breeding birds, forestry, monitoring, plantations, Fundy National Park, New Brunswick Numerous studies have related changes in the structure and quality of habitats in temperate forest to variations of the diversity and abundance of the birds present (Freedman et al. 1994; Hunter 1999). However, accurate predictions cannot yet be made of the effects of forest harvesting and plantation estab- lishment on avian communities. This is due to geo- graphical variations in bird species and communities, and in their responses to changes in habitat structure and quality caused by forestry practices. Studies in Europe and the southern United States have found that intensively managed conifer planta- tions support relatively small populations of few bird species (Moss 1978; Repenning and Labisky 1985; Donald et al. 1998). A study in New Brunswick, however, found smaller effects on birds (Parker et al. 1994). Other relevant studies have examined the effects of clear-cutting and other harvesting practices on birds and their communities, including work in Nova Scotia (Freedman et al. 1981; Morgan and Freedman 1986), central Ontario (James and Peck 1995), and in the central and northeastern U.S. (Crawford and Titterington 1979; Titterington et al. 1979; Niemi and Hanowski 1984). In this contribution we summarize the results of a study of birds breeding in conifer plantations in the vicinity of Fundy National Park (FNP), New Brunswick. We examined a chronosequence of 11 plantations up to 21 years old to describe the avian communities occurring during post-establishment succession. We also examined birds in natural, mixed-species forest of the type that has not been clear-cut and converted into the plantations, to exam- ine consequences of the conversion for avian species, guilds, and communities. Methods Study Area The study area (centered at 45.66° N and 65.10° W) is in the Greater Fundy Ecosystem (GFE) in southeastern New Brunswick. The GFE consists of Fundy National Park, a protected area of about 206 km*, plus its nearby surrounding area, which is a mosaic of forestry plantations, clear-cuts, residual stands of natural mixedwood forest, and agricultural land. The GFE has been the locale for various studies of environmental stressors that are potential threats to the ecological integrity of the region, with a focus on the effects of forestry practices (Woodley etal. 199n), *See Documents Cited section 475 476 Our avian study sites are located within the Fundy Plateau Ecodistrict of the Southern Uplands Ecoregion of the Acadian Forest Region (Loucks 1962: Rowe 1972). The climax forest of the Ecodistrict is dominated by mixed-species stands containing Red and White spruce (Picea rubens and P. glauca), Balsam Fir (Abies balsamea), Sugar and Red maple (Acer saccharum and A. rubrum), and Yellow, White, and Mountain birch (Betula alleghaniensis, B. papyrifera, and B. cordifolia). Extensive gap-phase disturbances, largely associated with natural irruptions of Spruce Budworm (Choristoneura fumiferana), have resulted in most mature forest in the region having a mixed-species composition, with patches of angiosperm trees and regenerating conifers. _ The landscape of the GFE has also been greatly affected by a range of anthropogenic stressors asso- ciated with forestry, agriculture, tourism, and trans- portation infrastructure. In particular, much of the natural forest outside of Fundy National Park has been clear-cut and converted into conifer plantations to be harvested on about a 40-year rotation. The most frequently planted species is Black Spruce (Picea mariana), which is less vulnerable to suffer- ing damage from Spruce Budworm than other spruces or Balsam Fir. Some plantations are stocked with other conifers, such as Jack Pine (Pinus banksiana), Larch (Larix laricina), and Norway Spruce (Picea abies). THE CANADIAN FIELD-NATURALIST pee | Vol. 116 Study Sites The five reference stands are representative of the natural, mixed-species forest that has not been har- vested and converted into nearby plantations. As defined here, angiosperm-dominated reference stands (RA) have >70% hardwood basal area; mixedwood stands (RM) have 36-70%, and conifer- dominated stands (RC) have <36% (Table 1). The reference stands were in a mature condition at the time of our study, although they had been selectively logged at various times prior to the establishment of the national park in 1948, when there were sawmills in the area and small settlements for which firewood was harvested (Burzynski 1985). The 11 plantations studied are located within 5 km of the park, and were chosen to represent the age range present when we began our study in 1993 (i.e., ranging from 3 to 21 years old). The plantations were planted to Black Spruce, except for one Larch stand. The plantations also had a vigorous natural regeneration of Balsam Fir, as well as various species of angiosperm shrubs and monocotyledonous and dicotyledonous forbs (Veinotte et al. 2002*). In all cases, the plantations had been established after the clear-cutting of natural, mature, mixed-species forest of a type similar to our reference stands. After they were clear-cut, the sites were mechanically site- prepared and then planted to conifer seedlings. The plantations received a herbicide treatment to suppress competing vegetation, intended to allow faster growth of the planted conifers. TABLE |. Description of the 16 study stands and sampling design. Reference stands (R) have prefix “A” if hardwood-domi- nated, “M7” if mixed-species, and “C” if conifer-dominated; plantations are denoted by “P” plus the age since establishment. Prominent tree species Age Area Year Site (% total basal area) (yr) (ha) Surveyed Reference Forest RAI As (26), Ba (21), Pr (17), Ar (17) 60 13.4 1992 RM1 As (35), Pr (27), Ar (18), Fg (16), Ba (4) 55 10.0 1993 RM2 Pr (59), Ba (27), Ar (8), Bc (6) 105 10.0 1993 RCI Pr (74), Be (11), Ar (10), Ba (3) 65 11.8 1992 RC2 Pr (67), Ba (16), Ar (13), Bc (3) 85 10.0 1993 Plantations P21 Pm* (86), Ab (12), Pg (1) 21 10.0 1993 P15 Pm* (78), Ab (22) 15 10.1 1992 P13 Pm* (82), Ab (13) 13 10.0 1993 P8 Pm* (61), Ab (38), Be (2) 8 10.0 1993 P7a No trees i | 14.1 1993 P7b Pm* (78), Ab (22) di 10.0 1993 P6 Ab (84), L1* (16) 6 10.3 1993 PSa No trees 5 10.0 1993 PSb No trees 5 10.0 1993 P4 No trees 4 10.0 1993 P3 No trees 3 Lis 1992 Species codes: Ab = Abies balsamea; Ar = Acer rubrum; As = Acer saccharum; Ba = Betula alleghaniensis; Bc = Betula cordifolia; Bp = Betula papyrifera; Fg = Fagus grandifolia; Pg = Picea glauca; Ll = Larix laricina; Pm = Picea mariana; Pr = Picea rubens; * = planted stock. 2001 JOHNSON AND FREEDMAN: BIRDS IN FORESTRY PLANTATIONS 477 All of the study sites are well-drained and located on flat terrain with gentle slopes less than about five degrees. The soil association is Lomond (Woodley 1985*), a humo-ferric podzol (Canada Soil Survey Committee 1978; Maliondo et al. 1990). All sites occur in a relatively small area, and are within 4 km of at least one other study site. Habitat Measurements Living and dead (i.e., snags) tree-sized stems (i.e., having a diameter at breast height (DBH) > 5 cm) in reference stands were sampled in 12 quadrats of 20m X 20m. Because the distribution and size spectrum of trees in plantations were relatively homogenous, they were sampled using fewer (9) quadrats of a smaller size (10m X 10m). Shrub- sized plants (DBH < 5 cm) were measured in two 5m X 5m quadrats nested in opposite corners of each tree quadrat. The length and diameter of coarse woody debris (CWD; diameter > 5 cm) were also recorded in the tree quadrats. The ground vegetation was sampled in 30 quadrats (1m X 1m) placed randomly along tran- sects running through each site. The percent cover of each plant species was visually estimated, and could exceed 100% due to layering of the foliage. The vertical distribution of plant cover was esti- mated at 25 evenly spaced points per stand. Vertical cover was estimated as the average cover of foliage obscuring a 5-cm-wide, 4-m-tall pole, within each of four height intervals (0-1 m, 1-2 m, 2—3 m, and 3-4 m) when viewed at a distance of 10 m in each of the four cardinal compass directions. Overhead canopy cover was estimated by sighting upwards through a 4.2-cm diameter, 10.5-cm long cylinder, from a height of 1.75 m. The height of the overhead canopy was also estimated. Bird Census Breeding birds were censused at 11 plantations and five reference sites (six during the 1992 field season, and an additional 10 in 1993). All plots were at least 10 hectares in area, and the spot-mapping technique was used to census birds (International Bird Census Committee 1970). Each plot was sur- veyed 10 times from the last week of May to mid- July, including two evening counts. Surveys were not conducted on windy or rainy days, as these con- ditions interfere with audibility. Territories were assigned on the basis of clusters of observations, coupled with delineation of territorial boundaries between individual males using records of simulta- neous singing (Bibby et al. 1992). Full or half-terri- tories were designated, with a “+” designation given in cases where a smaller part of a territory occurred within a census plot (these were assigned a value of 0.2 in the data analyses; Kirk et al. 1996). The data were standardized as the number of territories per 10 hectares. Species observed more than once on a cen- sus plot, but not having sufficient observations to be assigned part of a territory (as defined by Bibby et al. 1992), were identified as visitors, or “v.” Species recorded once were identified as transients, or “t.” Data Analysis Density and species richness were determined for breeding birds and for nesting and breeding guilds, which can be useful indicators of the effects of man- agement practices (Thingstad 1997). We examined five nesting guilds (ground, shrub canopy, tree canopy, variable canopy, cavity) and seven foraging guilds (aerial, bark, foliage, ground, nectar, raptor, tree driller). The Shannon-Weaver index was used as an indicator of species diversity (Shannon and Weaver 1949), where p; is the relative density of each species. Species similarity (S;) and population similarity (Sp) were calculated as indicators of the similarity of avian communities between sites (Legendre and Legendre 1983). The following habitat variables were calculated as averages among the sampling quadrats and used in the analyses: tree density (TDE; stems/ha); tree basal area (TBA; m?/ha); percent angiosperm tree basal area (THW; %); average tree diameter (TDI; cm); coefficient of variation of tree diameter (CVTDI; %); tree species richness (TSR); tree species diversity (TSD); shrub density (SDE; stems/ha); shrub basal area (SBA; m*/ha); percent angiosperm shrub basal area (SHW; %); shrub species richness (SSR); shrub species diversity (SSD); total stem density (TST, as sum of TDE and SDE; stems/ha); total basal area (TOB, as TBA + SBA; m7/ha); snag density (SND; stems/ha); snag basal area (SNB; m7/ha); volume of coarse woody debris (CWD; m°*/ha); total vertical cover (VCV, as sum of cover in four 1|-m intervals; %); percent shrub cover (SCV; average cover in three l-m intervals above the ground; %); total ground cover (HCV; sum of all species of ground vegetation; %); canopy cover (CCV; %); coefficient of variation of vertical cover (CVVCV; %); canopy height (HGT, m). Note that some of these variables were used as indicators of the horizontal and vertical patchiness of selected habitat variables (Roth 1976). Coefficient of variation was calculated as the sample standard devi- ation divided by the mean (Zar 1984). After viewing scatter plots of the densities of bird variables versus habitat variables, selected relation- ships were assessed using Pearson correlations. These correlations are intended for descriptive pur- poses, as rigorous use of this statistic is precluded by: the high within-stand variation of habitat vari- ables, the high number of zero values for some habitat and avian variables, and the relatively small number of stands being compared. Multivariate analyses were used to characterize avian “communities” (Gauch 1982). Hierarchical cluster analysis was used to group and classify the 478 sites by bird species densities (SYSTAT 1990). An average linkage cluster was used, using the unweighted pair-groups method, which calculates arithmetic averages (UPGMA; Sneath and Sokal 1973). The Pearson correlation was used for the sim- ilarity matrix in this cluster analysis (SYSTAT 1990). Two-way indicator species analysis (TWINSPAN; Hill 1979) was used to display the species in an ordered table, and facilitated classify- ing the bird species by habitat (Table 2). Ordination techniques were used to explore the relationships among stands. Although we have a relatively low number of stands in comparison to the number of variables, ordination is nevertheless useful in showing trends, if they can be explained ecologically (Gauch 1982). We used two types of ordinations: one based on bird species data, and the other on habitat data. The ordinations of stands were compared to see if bird communities and habitat variables showed similar relationships among the stands. Detrended correspondence analysis (DCA) was used to analyze the bird species data, and allowed the plotting of bird species in relation to the study stands. Indirect gradient analysis was per- formed by correlating habitat variables to the DCA axes, using Pearson correlation coefficients. Results and Discussion Bird Density A total of 72 bird species was recorded as breed- ing, foraging, or transient on at least one site (41 species had >1 territory/10 ha on at least one plot; Table 3). The total density of breeding birds ranged TABLE 2. Summary of bird community variables. Species diversity is based on species breeding on the plot; richness includes species breeding and foraging. Stand Bird Species Species Age Density Diversity Richness Stand (yr) (pr/10 ha) (H’) Plantation P3 3 16.5 e/a i P4 4 30.6 1.30 1] P5a 5 43.4 137 12 P5b 5 42.8 1.45 9 P6 6 54.3 1.88 18 P7a 7 48.3 Loh 20 P7b 7 56.2 1.41 12 P8 8 55.4 1.94 15 P13 13 84.6 2.13 2 P15 15 104.6 2.64 36 P21 21 86.0 2.59 24 Reference RM1 45 54.0 253 25 RAI 50 44.1 2.92 32 RC] 65 68.3 3.07 4] RC2 85 S11 2.48 27 RM2 95 70.3 2.84 a2 THE CANADIAN FIELD-NATURALIST Dy 3 Vol. 116 - from 16.5 pairs/10 ha in the 3-yr-old plantation to 104.6 pairs/10 ha in the 15-yr-old plantation (Table 2). The reference, mixedwood stands ranged from 44.1 to 70.3 pairs/10 ha. Bird density reached pre- cutting (i.e., reference) levels within 5 years of plan- tation establishment, and peaked at 15 years. The densities of breeding birds are similar to those reported in other studies of conifer plantations and mixedwood and coniferous forest in northeastern North America. DesGranges (1980) found 66.7 pairs/10 ha in a mixedwood stand in La Mauricie National Park, Quebec. Welsh and Fillman (1980) reported 102-197 pairs/10 ha in clear-cuts of conifer forest aged 11-24 years in northwestern Ontario. Christie (1993) found 18.5—47.2 pairs/10 ha in forest plots in Fundy National Park. Parker et al. (1994) found 33.3—-49.3 pairs/10 ha among stands of conifer forest and plantations up to 17-years old in south- eastern New Brunswick. Additional relevant studies include Erskine (1977) and James and Peck (1995). A peaking of avian density in mid-successional stands is a frequent observation in forest chronose- quences (Probst 1979; Morgan and Freedman 1986). However, Parker et al. (1994) reported a more linear relationship of avian density and stand age in a chronosequence of plantations up to 17 years old. Species Richness and Diversity Species richness (including species using the site for nesting or foraging) ranged from 9 to 36 in the plantations, and from 25 to 41 among reference stands (Table 2). By 13 years, species richness in the plantations was similar to that in reference stands. However, the richness of breeding species (1.e., hav- ing a density of “+” or more) in plantations ranged from 7 to 22 (mean + S.D., 11.8 + 4.9) compared with 18 to 31 in reference stands (24.2 + 5.1), a sig- nificant difference (ANOVA, p <0.001). Shannon- Weaver diversity (H') in reference stands ranged from 2.5 to 3.1. Plantations reached reference levels (H' > 2.5) after about 13 years (Table 2). The species diversity in younger plantations (3-8 years old) was 1.3 to 1.9, significant lower than in older plantations (2.1 to 2.6; p <0.001; ANOVA). The older plantations provided habitat for species charac- teristic of both younger and older stages of succes- sion, and this accounted for their relatively high diversity. Similarity Although the avian density, species richness, and species diversity reached levels similar to reference stands within 13 years, most of the bird species inhabiting the plantations were different from those in the natural forest. This observation is apparent in the matrices of species similarity (S,) and population similarity (S,) (Table 4). 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P2i° “RMI -RAI~ RCI P4 Forage P3 Code Species Ovenbird 0.8 2.8 Fa) Mourning Warbler [5:3 10.0 NO) 4.0 [0-922 8.0 Common Yellowthroat Wilson’s Warbler Canada Warbler Song Sparrow P| iO 60 59 102 id 215 [s,5 lei0 17.3 0 15.35 Lincoln’s Sparrow Swamp Sparrow 1.7 0.4 oD 4.0 126: 3:35 4.0" 795 18.0 8.4 1.0 1.0 3.0 35 White-throated Sparrow Dark-eyed Junco 4.0 2.0 THE CANADIAN FIELD-NATURALIST 0.4 Rose-breasted Grosbeak Pine Grosbeak Purple Finch 0.5 0.5 + 0.8 ie 2.0 PRFI Vol. 116 - tations, and vice-versa. Because the middle-aged plantations (i.e., P13, P15, and P21) supported some species of younger habitats as well as some of mature forest, these stands tended to have relatively high values of S, and S,. The differences between reference forest and plantations were particularly great for Sp, because of the large differences in populations, even if there were some species in common. Overall, populations varied less between sites than did species composition. Nesting and Foraging Guilds The avian community of younger plantations was strongly dominated by ground-nesting species, par- ticularly Common Yellowthroat (see Appendix 1 for binomials), Lincoln’s Sparrow, Song Sparrow, and White-throated Sparrow. Ground-nesting birds accounted for 86—100% of the avian density in plan- tations aged 3 to 8 years, decreasing to 38-55% in the oldest plantations studied (15 and 21 years post- establishment) and 21—27% in the reference stands. In contrast, canopy-nesting species such as Magnolia Warbler, Ruby-crowned Kinglet, Yellow-bellied Flycatcher, and Yellow-rumped Warbler were most prominent in older stands. They accounted for 63-76% of the avian density in reference stands and 44-61% in the oldest plantations, compared with 0-12% in plantations aged three to eight years (Alder Flycatcher, which nests in shrubs, was the first species in this guild to invade the plantations, at six years post-establishment). Cavity-nesting birds only occurred in significant numbers in reference forest, where the most promi- nent species were Black-capped Chickadee, Downy Woodpecker, Hairy Woodpecker, and Yellow-bel- lied Sapsucker. Although the plantations had few cavity-containing snags, any that did exist were used by such dependent species as Northern Flicker, Tree Swallow, American Kestrel, and Eastern Bluebird. The presence of natural cavities is well known as a limiting factor for these and other cavity-dependent species (Niemi and Hanowski 1984; Zarnowitz and Manuwal 1985; Adkisson 1988; Shreiber and Decalesta 1992; Freedman et al. 1996). In a separate study, artificially erected natural cedar cavities in plantations up to 17 years old were used by various species requiring this critical habitat element (including Boreal Chickadees, a forest species, in the 17-year-old stand; Freedman et al. 1996; Woodley et al., personal communication). The most rapid shift in nesting guilds occurred in the 13-15 year-old plantations, which provided habi- tat suitable for similar numbers of species of ground- and canopy-nesters. This transition occurred much sooner after the clear-cutting of hardwood forest in Nova Scotia (Morgan and Freedman 1986), where vegetative regeneration (by stump-sprouting) of cut trees and angiosperm seedlings restored a shrub- and tree-level canopy much more quickly than in the 2001 TABLE 4. Matrix of values of bird species similarity (Ss; left side of matrix) and population similarity (Sp; right side). RC2 0.14 Cals RM2 0.16 0.14 0.18 0.15 0.24 0.31 0.14 0.21 0.48 0.59 0.57 0.72 0.77 0.67 1.00 0.74 RMI 0.14 O7 0.11 RCI 0.24 0.15 0.19 0.16 0.37 0.39 nla 0.18 0.44 0.62 0.62 0.82 1.00 0.50 0.65 Us5 RAI 0.16 0.19 0.18 0.20 0.24 0.31 0.14 0.21 0.41 0.59 0.64 1.00 0.48 0.56 0.49 0.52 P15 P21 0.34 0.21 0.29 0.27 0.48 0.50 0.33 0.43 0.67 P4 P5a P5b P6 P7a P7b P8 Pi3 P3 S/S» P3 0.29 0.23 0.22 0.18 0.29 0.32 0.22 0.26 0.55 0.63 1.00 0.31 0.46 0.37 0.41 0.39 0.49 0.55 0.41 0.52 1.00 0.68 0.53 0.10 0.30 0.11 0.56 0.62 0.59 0.58 0.55 0.63 0.67 1.00 0.47 0.50 0.17 0.04 0.05 0.02 0.02 0.02 0.69 0.61 0.83 0.67 0.53 0.69 1.00 0.58 0.28 0.23 0.08 0.02 0.04 0.00 0.00 0.00 0.60 0.45 0.63 0.55 0.84 1.00 0.62 0.73 0.33 0.36 0.10 0.03 0.07 0.02 0.03 0.03 0.51 0.41 0.47 0.44 1.00 0.72 0.43 0.76 0.47 0.49 0.18 0.06 0.09 0.02 0.06 0.06 0.54 0.70 0.76 1.00 0.58 0.83 0.75 0.68 0.30 0.27 0.08 0.02 0.05 0.02 0.02 0.02 0.69 0.70 1.00 0.80 0.45 0.66 0.85 0.55 0.21 0.57 1.00 0.80 0.82 0.48 0.73 0.68 0.60 0.21 1.00 0.47 0.40 0.39 0.34 0.36 0.31 0.31 0.17 0.14 0.08 0).02 0.08 0.06 0.09 0.06 P4 0.10 0.06 0.13 0.17 0.05 0.14 0.37 0.54 0.63 0.71 0.71 0.69 Oi) 1.00 PSa P5b P6 0.06 0.09 0.13 0.05 (O} I) 0.31 0.39 0.57 0.74 0.70 1.00 0.53 0.57 P7a JOHNSON AND FREEDMAN: BIRDS IN FORESTRY PLANTATIONS 481 P7b P8 P13 P15 P21 1.00 0.49 Onis ‘OR Ps) 0.17 0.05 0.02 0.04 0.01 0.01 0.01 0.18 0.07 0.02 0.04 0.00 0.01 0.00 RAI RCI 0.55 0.36 0.62 0.46 0.15 0.29 0.23 RM] RM2 RC2 0.26 0.21 plantation chronosequence examined in the present study. Planted conifers have a relatively slow height growth compared with hardwood sprouts and seedlings, and plantation management includes the use of herbicide to release small conifers from the effects of competition. Most species at all sites were ground searchers or foliage gleaners. Ground searchers varied little across the plantation chronosequence, accounting for 10-34% of the avian density in stands aged 3-8 years, and 14—26% in those aged 13-21 years, com- pared with 18-27% in reference forest. Prominent species in this guild included White-throated Sparrow and Ovenbird. Foliage searchers also varied little among these habitat types, accounting for 61-90% of the avian density in plantations aged 3-8 years, 52-64% in those aged 13-21 years, and 58-70% in reference forest. Prominent species in this guild included Common Yellowthroat and Yellow-rumped Warbler. Other guilds were much less abundant. Species of the flycatcher guild were most abundant in planta- tions aged 6—21 years old, where their relative densi- ty ranged from 9 to 21%. Alder Flycatcher and Yellow-bellied Flycatcher were the most prominent species in this guild. Bark gleaners, such as Black- and-white Warbler, Red-breasted Nuthatch, and Brown Creeper, only occurred in reference forest, accounting for 2-7% of the avian density. Young plantations lack the foraging substrates for this guild (Franzreb and Ohmart 1978). Aerial foragers, such as Common Nighthawk and Tree Swallow, only occurred in plantations 8 years and younger, likely due to their preference for open nesting habitat (Ehrlich et al. 1988). Prominent Bird Species On average, six species accounted for 72% of the total avian density in plantations: Lincoln’s Sparrow, 20.4%; Song Sparrow, 13.7%; Common Yellowthroat, 12.8%; White-throated Sparrow, 12.7%; Yellow-bellied Flycatcher, 6.2%; and Magnolia Warbler, 5.4%. Seven mostly different species accounted for 57% of the density in reference forest: Black-throated Green Warbler, 11.2%; Blackburnian Warbler, 10.1%; Ovenbird, 8.3%; Magnolia Warbler, 7.9%; Yellow-rumped Warbler, 7.6%; Golden-crowned Kinglet, 7.0%; and Yellow-bellied Flycatcher, 5.3%. Habitat A synopsis of selected habitat variables is present- ed in Table 5 (more detailed data are in Johnson (1997*), Fleming and Freedman (1998), and Veinotte et al. (2002*)). The plantation data illus- trate the successional habitat changes occurring as the conifer forest develops. The average height of the canopy increased rapidly from about 1.1 m ina 3-year-old plantation to 6.9 m in the 21-year-old 482 THE CANADIAN FIELD-NATURALIST Vol. 116 plantation, compared with 10.6-16.6 m in reference stands (Table 5). Tree basal area increased steadily =NDAOh st ; : Syne 2 Fla sAdNSdoves in plantations, achieving 17 m*/ha in the 21-year-old MIHNKHnannnewo- A 2 5 stand, compared with 25-29 m*/ha in reference x Beith ss! diag, = stands. Trees in the plantations were predominantly O(2e-F % Snyee the planted conifers, along with natural regeneration of balsam fir and a few hardwoods that survived the g es. e herbicide treatment. The reference stands, in con- GIT eA oSevregn trast, are mixed-species communities of various coniferous and hardwood species (Table 1). Shrub Saint. Ee ne} lyhe & density was variable among the plantations, largely Siar S °5 06 eeitsc depending on the relative abundance of certain angiosperm species, particularly Sambucus race- = poh Be eS Bs mosa and Spiraea latifolia. a feet Seu ae Snags were scarce in the plantations, having been depleted during the clear-cutting and subsequent site BR ee 1 fs yee preparation. Snags were considerably more abundant Suite eye Se aca 2 in the reference forest (185—708/ha), partly because of mortality associated with past infestation with tal) ta Bis, each ey Spruce Budworm. Coarse woody debris was abun- ule Settee pale Sie dant in the younger plantations (48-52 m*/ha in the 3- and 4-year-old stands), due to the recent deposi- ch ee le a tion of logging slash. This deadwood component RIldiasasontonran became progressively depleted as the plantations aged (to 0.6 m°/ha in the 21-year-old stand). Coarse a od ean ey woody debris was relatively abundant in the refer- Bal eaiedien of Gqenve ey eren ence stands (13-45 m?/ha), due to the effects of tree mortality associated with Spruce Budworm. Further ae ee te gh details on deadwood components in this chronose- BISSOSRASONOAAN quence, including projections to plantation maturity and over several rotations, are in Fleming and 3S Saco ti) © Freedman (1998). Both snags and coarse woody i or So. one S ; : : ; Sl kak a ak al debris are crucial elements of the habitat for a wide range of bird species and numerous other compo- Solan oe ona S nents of forest biodiversity (Newton 1994; Freedman SHARISDSONYTOMOMHS = et al. 1996). - The cover of ground vegetation increased rapidly m2 oe = during the first few years of plantation development s le) 5 a) (=> Laer x p p > Seieeuege 8e ae SS leveling off at about 100—140% in stands older than a 2, six years. The cover of ground vegetation was less in Sis Bie he ref d ing from 55-98%, d el ars CRN eee a the reference stands, ranging from 55—98%, due to Soueoe ts hh gs the relatively dense overhead canopy. Plant species cs of the youngest plantations were predominantly rud- 3 = lg eee aA as eral forbs (e.g., Aster spp., Epilobium angustifolium, $ See ea) cite Maes and Solidago spp.) and graminoids (e.g., Agrostis 5 a spp. and Scircus cyperinus) that invaded the dis- 3 oO] & o6 a turbed site, plus an in-situ regeneration of plants that 3 survived the clear-cutting and site preparation (e.g., 3 g <2 Cornus canadensis, Dennstaedtia punctilobula), and 9 ae 5 others that germinated from a persistent seedbank ro) sé z aRe é (e.g. Rubus strigosus and Sambucus racemosa). o Set easue SE Details on species composition of the ground vegeta- E 21s S oe = SS 5s = tion in this chronosequence are in Veinotte (1998*) iano Al = Al E|z22= 28 2 a ® “ob and Veinotte et al. (2002*). WHelasases2goq0sg 22 - sauvunyses > & e : rf s|35 Soo g ou > Multivariate Analyses ais ssgee a 3 38 5 2 A cluster analysis was used to analyze the matrix FIZTIFPRPAHHORKDOO of 41 bird species densities by 16 stands (Figure 1). 2001 JOHNSON AND FREEDMAN: BIRDS IN FORESTRY PLANTATIONS 483 RM1 OLD2 | 233 FIGURE 1. Dendrogram showing the results of the cluster analysis of 16 sites based on densities of 41 bird species. The first division separated stands 15 years old and younger from older ones. A second division resulted in a total of four distinct clusters: ¢ YNGI consists of the youngest plantations, aged 3 to 8 years. Breeding birds common to all stands in YNGI1 include Common Yellowthroat, Song Sparrow, Lincoln’s Sparrow, and White-throated Sparrow. Dark-eyed Junco was present at all but one site. This group is comprised of ground-nesting, open-canopy species. ¢ YNG2 includes intermediate-aged plantations aged 13 and 15 years. Birds occurring in both sites include Yellow-bellied Flycatcher, Ruby-crowned Kinglet, Hermit Thrush, Magnolia Warbler, Yellow-rumped Warbler, Palm Warbler, Common Yellowthroat, Lincoln’s Sparrow, White-throated Sparrow, and Dark-eyed Junco. These species are a mixture of ground-nesting and forest-nesting species capable of inhabiting young, second-growth coniferous forest. ¢ OLDI1 consists of the most conifer-dominated refer- ence stands (RM2, RC1, and RC2) plus the oldest plantation (P21). Birds occurring in all stands of this group include Yellow-bellied Flycatcher, Golden- crowned Kinglet, Ruby-crowned Kinglet, Swainson’s Thrush, Magnolia Warbler, Yellow- rumped Warbler, Black-throated Green Warbler, Blackburnian Warbler, Bay-breasted Warbler, and Dark-eyed Junco. These are mostly canopy-nesting species of coniferous or conifer-dominated mixed- wood forest. * OLD2 consists of the most angiosperm-rich refer- ence stands (RAI and RM1). Birds common to this cluster include Yellow-bellied Sapsucker, Golden- crowned Kinglet, Swainson’s Thrush, Hermit Thrush, Blue-headed Vireo, Red-eyed Vireo, Black- throated Blue Warbler, Yellow-rumped Warbler, Black-throated Green Warbler, Blackburnian Warbler, Bay-breasted Warbler, Ovenbird, Dark- eyed Junco, and Purple Finch. These species are either typical of mature hardwood forest, or of hard- wood-dominated mixedwood. Most are canopy-nest- ing species, except for three ground nesters (Hermit Thrush, Ovenbird, and Dark-eyed Junco). The data matrix of bird density for 16 stands was also analyzed by detrended correspondence analysis (DCA). A subset of the 41 most abundant species was used, excluding rare species that could have a disproportionate effect on the results of the DCA. Indirect gradient analysis was then performed by correlating habitat variables with the DCA axes. The first axis of the DCA explained 43% of the variance, and was negatively correlated to habitat variables that had their highest values in reference stands (Table 6). This axis separates the reference stands from the young plantations, with older plantations in between (Figure 2). Although the data are not pre- sented here, the ordination of bird species on Axis 1 clearly separates species of mature reference forest from those of younger plantations. The second DCA axis explains much less of the variation (5.2%) and has relatively weak associations with habitat vari- ables. It is most strongly related to high shrub basal area and stem density, as well as high total cover of ground vegetation. Both the cluster analysis and the ordination sup- port the following observations: (a) the avian “com- munities” of younger plantations are dissimilar to those of both reference forest and older plantations, (b) as plantations age they become somewhat similar to reference forest in their avian community, and (c) like-aged stands support similar avian communities. However, the oldest plantations that we studied (15 and 21 years post-establishment), which are about 1/3 to 1/2 of their harvest rotation, are quite different from the natural, reference forest in terms of the structure and complexity of their habitat and vegeta- tion, and to a lesser degree in their avian community. General Discussion Clear-cutting and the subsequent establishment of conifer plantations result in temporary habitat oppor- tunities for open-canopy species of birds, such as Alder Flycatcher, Common Yellowthroat, Song Sparrow, and Lincoln’s Sparrow. Interestingly, some of these species are relatively uncommon in Fundy National Park. The total populations of Song Sparrow and Lincoln’s Sparrow in 1992 were esti- mated as fewer than 100 pairs each, and that of Alder Flycatcher about 130 pairs (Christie 1993*). 0 100 200 300 400 500 DCA1 FiGuRE 2. Study sites displayed on the first two axes of a Detrended Correspondence Analysis performed using densities of 41 bird species in 16 study sites. ee 484 TABLE 6. Correlations between habitat variables and the first two axes of a detrended correspondence analysis per- formed using the densities of the 41 most abundant bird species in 16 study stands. Axis 1 accounted for 43.4% of the variance, and axis 2 5.2%. See Methods section for explanation of habitat acronyms. Habitat Variables DCA Axis 1 DCA Axis 2 TBA -0.993 -0.036 TOB -0.990 0.037 CVTDI -0.968 -0.067 TDI -0.945 0.024 HGT -0.938 -0.087 TSR -0.903 0.160 CCV -0.870 -0.160 AGE -0.862 -0.050 THW -0.818 -0.026 TSD -0.807 0.205 SCV _ -0.739 0.300 SNB -0.692 -0.118 VCV -0.688 0.335 SND -0.653 -0.091 SHW -0.571 -0.021 TDE -0.549 ‘0.146 SBA 0.070 0.553 HCV 0.105 0.521 TST -0.241 0.501 SDE -0.163 0.499 SSR -0.082 0.457 r= 0.742; p= 0.001 p=625:p—0.01 N= 16, di=14 In addition, Common Nighthawk may not be breed- ing in the national park (Christie 1993*), but it uti- lizes habitat in nearby clear-cuts and young planta- tions. Silvicultural practices such as thinning and herbicide use help to maintain plantations in a rela- tively open condition for an extended period of early-successional time, perhaps prolonging their suitability for some of these species (MacKinnon and Freedman 1993). As the plantations mature they provide habitat for some species that inhabit conifer-dominated forest, such as Yellow-bellied Flycatcher, Ruby-crowned Kinglet, Nashville Warbler, Magnolia Warbler, and Yellow-rumped Warbler. We did not have the opportunity to examine plantations older than 21 years (because none existed in our study area). However, based on observations of natural conifer- dominated forest and our oldest plantation, we pre- dict that plantations older than about 25 years would also provide habitat suitable for Golden-crowned Kinglet, Blackburnian Warbler, Bay-breasted Warbler, Black-throated Green Warbler, and Cape May Warbler, as well as some other species. In addi- tion, if older plantations are adjacent to natural forest, they are utilized as foraging habitat by some forest species, such as Black-capped Chickadee, Boreal Chickadee, and Red-breasted Nuthatch. THE CANADIAN FIELD-NATURALIST Vol. 116 However, a total of 16 species that bred in natural reference forest were not observed in plantations, including species that prefer or require mature habitat with a significant component of angiosperm trees, and those that require cavities. In our study area, species that will probably become at risk from the extensive conversion of hardwood-dominated or mixedwood stands into coniferous plantations include Ovenbird, Red-eyed Vireo, and Black-throated Blue Warbler (see also Finch 1991; Baguette et al. 1994; Donald et al. 1998; Easton and Martin 1998). As noted previ- ously, all cavity nesters are at risk from plantation management of the type we studied, which results in the destruction of all or most snags during clear-cut- ting and site preparation. Moreover, there are poor prospects for the development of new cavity trees during subsequent plantation rotations (Fleming and Freedman 1998). Proposed Mitigations Although our research was not specifically designed to test hypotheses directly related to habitat mitigations, we believe that we can make several useful recommendations based on the results of this study. We stress, however, that these recommenda- tions should be subjected to field tests to ensure that they are efficacious. In any forest-management area, reasonably sized patches of all naturally occurring forest types should be protected from conversion into plantations or other kinds of anthropogenic ecosystems. We cannot make specific suggestions about the sizes of the protected areas, but they should be large enough to be self- maintaining over the longer term. If this is done, then most indigenous species of birds, and other elements of native biodiversity, should be able to maintain viable populations in the management area. In our study area (the Greater Fundy Ecosystem), we recommend that forest managers should ensure that adequate areas of closed-canopy, mixedwood, and angiosperm-dominated forest remain available to pro- vide habitat for species for which plantations appear to be unacceptable. In part, this objective could be achieved by allowing some clear-cuts and other har- vested areas to regenerate naturally after logging. Areas converted to plantations should be allowed to retain or regenerate a substantial component of deciduous trees. In part, this could be achieved by deliberately leaving some areas untreated during herbicide sprays. In fact, about 10% of treated areas are “missed” during typical silvicultural herbicide applications (Santillo et al. 1989; MacKinnon and Freedman 1993). We recommend that the non- sprayed area should be at least twice as large as this, to ensure that the overall habitat retains sufficient _ islands of hardwood shrubs and trees for dependent species. Ensuring the persistence of snags, cavity trees, and coarse woody debris is also necessary to oe | 2001 JOHNSON AND FREEDMAN: BIRDS IN FORESTRY PLANTATIONS 485 maintain populations of species dependent on these critical habitat elements. Woodley and Forbes (1997*) have recommended that 10-12 snags (>20 cm DBH) plus 12-15 live mature aspen (Populus spp.) or beech (Fagus grandifolia) be retained per hectare, preferable in clumps of at least 25 m’, to provide these habitat elements in clear-cuts and plan- tations in the Greater Fundy Ecosystem. It is also necessary to ensure that new cavity trees develop during subsequent plantation rotations. We believe that this could result if “islands” of uncut, mature, hardwood-dominated forest are retained within clear-cuts and plantations. We recommend that these tree-islands should be at least 1 hectare in area, and should be present in at least a 1:10 ratio to area of clear-cut or plantation. Although the ecological integrity of these tree-islands will be challenged by their small area, high edge ratio, and silvicultural practices occurring around them, they should prove capable of maintaining a sufficient number of cavity trees, snags, and coarse-woody debris to maintain populations of at least some dependent species with- in the management block (Jansen et al. 1995). To some degree, riparian buffer strips adjacent to clear- cuts and plantations may provide these habitat ele- ments, although this service would be degraded if these linear features are selectively harvested of their larger trees. More detailed recommendations about deadwood elements of habitat in the GFE are in Woodley and Forbes (1997). Acknowledgments We are grateful to Fulton Lavender for his assistance throughout this project. We also thank Jill Adams, Tracy Fleming, Jonathan Freedman, Wolfgang Maass, Minga O’Brien, Kathleen O’Sullivan, Cindy Staicer, Cam Veinotte, and Ruth Waldick for assistance in the field and other support. We also thank wardens, biologists, and administra- tive staff of Fundy National Park for their assistance in various ways. J. D. Irving Forest Products Limited allowed access to their plantations, and provided information regarding management practices. This study was funded by grants to B. F. from the Natural Sciences and Engineering Research Council of Canada and the Greater Fundy Ecosystem Research Group, and by contracts from the Fundy Model Forest and the Canadian Wildlife Service University Research Support Fund. Accommodations during fieldwork were provided by Fundy National Park. A Dalhousie University Graduate Fellowship to Greg Johnson is also greatly appreciated. Documents Cited (marked* in text) Christie, D. S. 1993. Survey of breeding birds in Fundy National Park, 1992: Survey Methods and Results. Unpublished Research Report, Fundy National Park, Alma, New Brunswick. Johnson, G. A. M. 1997. The effects of forestry on breed- ing bird communities in the vicinity of Fundy National Park, New Brunswick. M.Sc. thesis, Department of Biology, Dalhousie University, Halifax, Nova Scotia. Veinotte, C. 1998. A comparative analysis of plant com- munities in natural, mixed-species forests and silvicul- tural plantations within the greater Fundy ecosystem, New Brunswick. M.Sc. thesis, Department of Biology, Dalhousie University, Halifax, Nova Scotia. Veinotte, C., B. Freedman, W. Maass, and F. Kirstein. 2002. Comparison of the ground vegetation in spruce plantations and natural, mixed-species forest in the Greater Fundy Ecosystem, New Brunswick, Canada. Submitted manuscript. Woodley, S. J. 1985. Fundy National Park, resource description and analysis. Parks Canada, Halifax, Nova Scotia. Woodley, S., and G. 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SYSTAT for DOS, Version 5.0. SYS- TAT, Inc., Evanston, Illinois. Thingstad, P. G. 1997. Challenges to conservation of biological diversity in boreal forestry landscape; A case study using bird guilds as environmental indicators. Fauna Norvegica (Series C) 20: 49-68. Titterington, R. W., H. S. Crawford, and B. N. Burgason. 1979. Songbird responses to commercial clearcutting in Maine spruce-fir forests. Journal of Wildlife Manage- ment 43: 602-609. Welsh, D., and D. R. Fillman. 1980. The impact of forest cutting on boreal bird populations. American Birds 34: 84-94. Zar, J. E. 1984. Biostatistical Analysis. Prentice-Hall, Inglewood Heights, New Jersey. 2001 JOHNSON AND FREEDMAN: BIRDS IN FORESTRY PLANTATIONS Zarnowitz, J. E., and D. A. Manuwal. 1985. The effects of forest management on cavity-nesting birds in north- western Washington. Journal of Wildlife Management 49: 255-263. Appendix 1. Scientific names of bird species. Ruffed Grouse, Bonasa umbellus Killdeer, Charadrius vociferus Common Snipe, Gallinago gallinago American Woodcock, Scolopax minor American Kestrel, Falco sparverius Common Nighthawk, Chordeiles minor Ruby-Throated Hummingbird, Archilochus colubris Yellow-Bellied Sapsucker, Sphyrapicus varius Hairy Woodpecker, Picoides villosus Downy Woodpecker, Picoides pubescens Northern Flicker, Colaptes auratus Eastern Wood-Pewee, Contopus virens Yellow-Bellied Flycatcher, Empidonax flaviventris Alder Flycatcher, Empidonax alnorum American Robin, Turdus migratorius Blue-headed Vireo, Vireo solitarius Tree Swallow, Tachycineta bicolor Black-Capped Chickadee, Parus atricapillus Boreal Chickadee, Parus hudsonicus Red-Breasted Nuthatch, Sitta canadensis Brown Creeper, Certhia americana Winter Wren, Troglodytes troglodytes Golden-Crowned Kinglet, Regulus satrapa Ruby-Crowned Kinglet, Regulus calendula Eastern Bluebird, Sialia sialis Swainson’s Thrush, Catharus ustulatus Hermit Thrush, Catharus guttatus Received 15 August 2000 Accepted 19 November 2002 Red-Eyed Vireo, Vireo olivaceus Nashville Warbler, Vermivora ruficapilla Northern Parula, Parula americana Magnolia Warbler, Dendroica magnolia Cape May Warbler, Dendroica tigrina Black-Throated Blue Warbler, Dendroica caerulescens Yellow-Rumped Warbler, Dendroica coronata Black-Throated Green Warbler, Dendroica virens Blackburnian Warbler, Dendroica fusca Palm Warbler, Dendroica palmarum Bay-Breasted Warbler, Dendroica castanea Blackpoll Warbler, Dendroica striata Black-and-White Warbler, Mniotilta varia American Redstart, Setophaga ruticilla Ovenbird, Seiurus aurocapillus Mourning Warbler, Oporornis philadelphia Common Yellowthroat, Geothlypis trichas Wilson’s Warbler, Wilsonia pusilla Canada Warbler, Wilsonia canadensis Song Sparrow, Melospiza melodia Lincoln’s Sparrow, Melospiza lincolnii Swamp Sparrow, Melospiza georgiana White-Throated Sparrow, Zonotrichia albicollis Dark-Eyed Junco, Junco hyemalis Rose-breasted Grosbeak, Pheucticus ludovicianus Pine Grosbeak, Pinicola enucleator Purple Finch, Carpodacus purpureus a ee ma ee ee a Testing a Double-Count Aerial Survey Technique for White-Tailed Deer, Odocoileus virginianus, in Québec FRANCOIS PotvIN!, LAURIER BRETON’, AND LOUIS-PAUL RIVEST? Société de la faune et des parcs du Québec, 675 boul. René-Lévesque est (lle), Box 92, Québec (Québec) GIR 5V7 Canada; email: FPotvin@fapaq.gouv.qc.ca 2Société de la faune et des parcs du Québec, 675 boul. René-Lévesque est (lle), Box 92, Québec (Québec) GIR 5V7 Canada 3Département de mathématiques et de statistique, Université Laval, Québec (Québec) G1K 7P4 Canada Potvin, Francois, Laurier Breton, and Louis-Paul Rivest. 2002. Testing a double-count aerial survey technique for White-tailed Deer, Odocoileus virginianus, in Québec. Canadian Field-Naturalist 116(3):488—496. In a double-count aerial survey, two independent observers, located on the same side of an aircraft, simultaneously count animals in sample plots. To evaluate if this technique could be implemented as part of our White-Tailed Deer (Odocoileus virginianus) management program, we assessed its precision to estimate densities over large wintering areas (2 25 km”, 10 surveys) and whole hunting zones (1600-26 000 km’, 14 surveys). We also tested its repeatability by replicating eight surveys two to five times. We finally compared double-count aerial surveys with pellet-group counts, which were previous- ly used to estimate deer numbers. Surveys of large wintering areas indicated that a 90% confidence interval (CI) of + 20% could be obtained with a sample size of 50-100 plots (5 km X 60 m strip plots). In hunting zones, 100—200 plots would have been needed to reach the same precision. Densities from replicated surveys were not considered biologically different (difference > 30% between two replicates within each survey) for 21 of 24 replicates overall. When both techniques were applied to the same wintering areas, the 90% confidence limits of the aerial estimate encompassed the pellet-group estimate in five of nine surveys, and was lower in one survey and was higher in three surveys. Although the costs of the aerial survey and the pellet-group count techniques are rather similar, we suggest that aerial surveys provide better estimates. We conclude that the double-count technique is reliable to survey White-Tailed Deer at a reasonable cost. In our context, a typical zone (200 plots) requires 30-40 helicopter hours and 10 days of work by a three-person crew. Key Words: White-Tailed Deer, Odocoileus virginianus, aerial survey, double count, Petersen estimate. Aerial surveys are commonly used to estimate the size of large mammal populations, notably Caribou (Rangifer tarandus), Elk (Cervus elaphus), and Moose (Alces alces) (Courtois et al. 1994; Couturier et al. 1996; Gasaway and Dubois 1987; Eberhardt et al. 1998). Such surveys are less common for White- Tailed Deer (Odocoileus virginianus) and Mule Deer (O. hemionus), as most jurisdictions rely on harvest statistics to manage populations. Pellet-group counts (Neff 1968) have historically been used to estimate deer numbers. Aerial surveys for deer have been conducted in several vegetation types: brush-domi- nated habitats in Texas (Beasom 1979; Beasom et al. 1981), Pinion Pine (Pinus edulis) — Juniper (Juniperus osteosperma) in Colorado (Kufeld et al. 1980; Bartmann et al. 1986, 1987; White et al. 1989), and deciduous forest in Connecticut (Kilpatrick and Ellingwood 1993), Missouri (Beringer et al. 1998) and Ohio (Stoll et al. 1991). This technique is difficult to apply in coniferous or mixed conifer-deciduous forests because of the closed tree canopy. Thermal imagery appears as a promising technique, particularly in deciduous for- est, but is not yet feasible under coniferous cover (Gill et al. 1997; Havens and Sharp 1998). Using correction factors in aerial surveys might be an alternative in coniferous forest landscapes. Correction factors should be specific to each survey, as many authors (Pollock and Kendall 1987; Graham and Bell 1989; Marsh and Sinclair 1989; Seber 1992) pointed out that it might be biased to use a correction factor obtained in one area under one set of conditions for another area or under different con- ditions. To evaluate and correct the sighting proba- bility, Floyd et al. (1979) and Beringer et al. (1998) have used radio-collared deer, a costly approach. Hence, a technique incorporating survey-specific correction factors in its application at a low cost is needed. The double-count technique provides such a survey-specific estimate of the sighting probability. This method relies on two observers searching inde- pendently all the parcels sampled and reporting their findings to a third party so that animals seen in com- mon or detected only by one observer can be tallied. Correction factors can be estimated with the Petersen model (Seber 1982; Pollock and Kendall 1987). We tested the double-count technique for White- Tailed Deer on Anticosti Island (Potvin et al. 1992; Rivest et al 1995) and in different areas on the Québec mainland. Our objective was to evaluate if this technique could be implemented as part of our provincial deer management program, based on (1) precision and repeatability of the density estimates under different conditions and (2) comparison of ’ deer number estimates and costs with the pellet- group count, the technique that was previously used. 488 2002 Study areas The double-count aerial survey technique was tested in Québec in eight large deer wintering areas (25-213 km7?), two forest blocks (160 and 475 km?) on Anticosti Island (zone 20), and 11 hunting zones (1 600-26 000 km?) on the mainland (Figure 1). Deer continuous distribution in the province is limit- ed to those zones. Southern zones (4, 5, 6, 7, 8) have a mixed agriculture-forested landscape while north- ern zones (1, 2, 3, 9, 10, 11, 20) are largely forested. Vegetation types are mostly deciduous or mixed, except on Anticosti Island which is dominated by boreal forest. During the snow season, deer congre- gate in wintering areas with a dense mixed and coniferous canopy. Winters are milder in the south, with three to four months of snow cover, but severe in the northern and eastern parts of the deer range, with almost six months of snow. Snow depth rarely exceeds 50 cm in the southern zones but can do so for 30 to 70 days on average in northern ones. Methods Aerial survey technique A detailed description of our double-count aerial survey technique can be found in Breton and Potvin (1997). Deer were counted by two observers sitting on the left side of a Bell 206-B (Anticosti) or Bell 206-L (other zones) helicopter. A bubble window installed in the back door enabled the rear observer POTVIN, BRETON, AND RIVEST: DOUBLE-COUNT AERIAL SURVEY 489 to see under and beside the aircraft. The front observer used the front upper, side and floor win- dows. The width of the narrow strip plot extended on the left side only from 0° (below the helicopter) to a 45° angle from the vertical. For each observer, the outer boundary was delimited by two reference marks denoted by plastic tape, on the side window and on the bottom of a rod extending outside and perpendicular to the aircraft. Before each survey, the helicopter followed a straight line (road, air strip) so that both observers verified that their viewing angles were identical. The navigator sat in the back seat behind the pilot and used a bubble window to provide front and later- al vision. He was responsible for discriminating and recording the deer groups seen. To ensure indepen- dence between the observers, the front observer, navigator, and pilot were connected to the communi- cation system of the aircraft and the rear observer was in contact with the navigator through a portable system. The earphones of the navigator were modi- fied in such a way that one ear was connected to the helicopter system and the other to the portable one. A switch enabled the navigator to speak separately to either observer. Surveys were conducted in winter. The survey area was delineated on 1:50 000 topographical maps. Large wintering areas were covered by systematic sampling and a 1—km buffer was usually added to a, 1 NN F \ 2 apa A Simone Saumon Armstrong Watopeka Lac David La Macaza Saint- Gédéon Island Brook Pohénégamook J Témiscouata — fa FicurE 1. Location of hunting zones (1-20), forest blocks (A, B) and deer wintering areas (C-J) where the double-count aerial survey technique was applied in Québec. 490 their periphery to include deer that might be outside of the traditionnal boundary of the yard. For hunting zones, three different sampling strategies were used depending on deer distribution: (1) systematic sam- pling of all forested areas in zones where deer were uniformly distributed and at high density (zone strat- egy), (2) systematic sampling with survey limited to wintering areas > 2.5 km? in zones with sparse distri- bution (yards strategy), and (3) stratified sampling with survey of both forested and wintering areas, for zones in between (mixed strategy). Strip plots 5—km long (6 km in earlier tests on Anticosti Island) were systematically distributed on the survey area along parallel lines oriented along a north-south azimuth. The same distance usually separated plots and lines to provide a complete systematic survey plan. Non- forested areas (lakes, agricultural and suburban areas) occupying > 300 m along the line were excluded as they are not deer habitat. Therefore, sur- vey plots were often discontinuous and made up of segments, especially in agro-forested landscapes. At the periphery of the survey blocks, incomplete plots on a line extended to the next line to cumulate 5 km. Loran-C (earlier surveys) or GPS navigation sys- tems of the aircraft were used to keep the bearing (Boer et al. 1989; Leptich et al. 1994). The pilot used the longitude reading to stay on line or to change lines (Beringer et al. 1998). The navigator deter- mined the beginning and the end of plots along lines using visual features or latitude readings. The pilot kept an altitude of 60 m, verified by a radar altime- ter, and a speed of 70 to 100 km/hr. Observers counted deer groups and classified them according to size (1, 2, 3, etc.) and activity (moving or inactive [bedded or standing immobile]). The navi- gator recorded deer sightings directly on the map and separately for each observer. Observers reported groups at the moment they were perpendicular to the helicopter, even if they had been detected in advance. Deer that had moved outside the strip before the pas- sage of the helicopter were recorded if they were inside when initially seen. Groups located outside the strip were not tallied. When observers counted a dif- ferent number of deer in a group, the higher value was used as group size in the computations. Computation of sighting probabilities and deer densities Because of the 45° viewing angle, the width of the strip is equal to the altitude (60 m), giving an area of 0.3 km? per plot (5 km X 60 m). A study area con- tains M plots where m of these are surveyed so that f = m/M is the sampling fraction. Let h = 1, ..., H, and i = 1, ..., m denote the subscripts representing the group size and the parcel sampled, respectively. The following notation is used: Nin, = Number of groups of h animals in parcel i seen by front observer only (k = 1), rear observer only (k = 2) or both observers (k = 3), THE CANADIAN FIELD-NATURALIST Vol. 116 nN), = Sum of the n,,,’s for m parcels sampled, n;,, = total number of groups of h animals seen in parcel 1 (nin, = Ning + Ning + Nin3)> n,, = total number of groups of / animals seen in m parcels surveyed. For sighting probabilities, we consider four group sizes: single deer, groups of two, groups of three, and groups of four or more. Sighting probabilities by observer (p,,) are computed according to the Petersen estimate (Magnuson et al. 1978) for each group size: N 43 N 3 Pi ag err and Poe (1 pp + 43) (n,,+n n3) The total population of deer and its variance are esti- mated using equations proposed by Rivest et al. (1995). These estimates use three correction factors (c,,), one for single deer, one for groups of two deer, and one for groups of three deer: Nal py gS eee n,(n,3+ 1) For groups of four deer or more, we assume a correc- tion factor of 1 (no correction). Based on 25 surveys, the sighting probability for such groups was on aver- age 0.92 and 0.90 for the front and rear observer, respectively (Figure 2). Since at least one observer had a sighting probability of 1.00 in 17 of these sur- veys, the correction factor c, = 1 most of the time. The variance estimator for c;, is computed as follows: (c, - 1) le, @,, - 2,3 - 2) +1) ae Fe See aaa An estimate of the total number of deer in each par- cel (N;), its variance and deer population in the sur- vey area (T) is given by: O FRONT OBSERVERG REAR OBSERVER 1.00 0.90 0.80 0.70 0.60 0.50 0.40 0.30 GROUP SIZE FIGURE 2 Sighting probability (+ 1 SE) of the front and rear observers according to group size obtained in 25 double-count aerial surveys in Québec. 2002 POTVIN, BRETON, AND RIVEST: DOUBLE-COUNT AERIAL SURVEY 491 H = CN + 2C2Nj2 + 3¢3n;3 + Lhn,, ; h=4 N. I var (N;) = [1/(m-1)] [YNi?- CON], pal frm (11) > N, . i=1 The variance of T has three components due to the variability of the selection of m parcels (0,2), the estimation of the correction factors (o,2), and the prediction of the number of deer (o,,”): [If (Uf) M varN; | - (1-f)o,’, (1/f) [n}.var(c,) + 4n>, var(c, ) + 9n3.var(c3)] , "Pi 2: Oo. 0,2 = (I/F?) {nj [e,? - ¢ - v(e)] + 4 ny [ep? - € - v(c>)] + 9 n3[c37 - c3 -v(c3)]} , wat 4'\0;? + 0,? . This variance is slightly larger than that presented by Rivest et al. (1995: Table 1) because some correction terms have been omitted to simplify formula expres- sion. To allow comparison among study areas, results are generally presented in terms of density ( 250 plots). Nonetheless, it was not pos- sible to evaluate the global precision of the deer popu- lation estimate because the variability of the other variables (defecation rate, SIZE, and PERIOD) is unknown. This precluded the use of a statistical test between the aerial survey and pellet-group count deer estimates. Therefore, we considered that both esti- mates were similar if the 90% confidence limits of the aerial estimate encompassed the estimate of the pellet- group count. We also compared the cost of both tech- niques using figures obtained from our surveys. Results Precision and repeatability In wintering areas, the 90% CI in 10 surveys var- ied from + 10 to + 29% (Table 1). To reach a + 20% precision, sample sizes of 50 to 100 plots would suf- fice. Precision was more variable in hunting zones, ranging from + 13 to + 49% (Table 2). In general, zones where sampling was limited to wintering areas provided narrower CI than those where all forested areas were surveyed. In most hunting zones, 100 to 200 plots would be necessary to achieve a 90% CI of + 20%. Assessment of repeatability was based on eight THE CANADIAN FIELD-NATURALIST UU ee a ne Vol. 116 surveys that were replicated two to five times, totalling 24 replicates (Table 3). The time delay between replicates of a specific survey ranged between 2 and 86 days. Deer densities 90% CI over- lapped among replicates within any survey. Three surveys each had a single replicate that had a differ- ence > 30% in density with other replicates of the same survey. Comparison with the pellet-group count technique Confidence limits for aerial survey deer estimates encompassed pellet-group count estimates in five of nine surveys of wintering areas (Table 4). Estimates from aerial survey were lower in one survey and higher in three surveys. Pellet-group counts take approximately 40 man- days to survey a single large wintering area (2 25 km7). Based on $200 CDN / person-day for salaries and $125 / day for travel expenses, this amounts to $13 000. A similar survey for a whole hunting zone costs about $32 500. The aerial survey of a large wintering area necessitates about 60 plots, which amounts to $8 600 for the helicopter ($1000/hr X 60 plots/7 plots per hr) and six person-days, for a total of $10 500. Surveying a hunting zone amounts to $33 300 for the helicopter ($1000/hr X 200 plots/6 plots per hr) and 30 person-days, for a total of $43 000. Discussion The double-count technique provided precise and repeatable estimates of White-Tailed Deer density at a reasonable cost. Sample size needed to achieve a 90% CI of + 20% for either wintering areas (50 to 100 plots) or hunting zones (100 to 200 plots) was relatively small. Much smaller plots (5 km X 60 m) TABLE 2. Precision obtained and sample size required for a 90% CI of + 20% in 14 aerial surveys of hunting zones using the double-count technique to estimate deer density in Québec. Sampling Survey strategy* (zone - year) m plots (x°) A 5 - 1990 61 of He 170 125 6 - 1990 82 i 8 - 1992 63 oi B 1 - 1993 89 0.03 2 - 1993 196 0.3 3 - 1996 201 1.6 9 - 1995 277 | 11 - 1992 184 2.0 C 4-199] 190 aes 4 - 1995 S22 a 6 - 1992 147 7.4 7 - 1996 199 oT 10 - 1994 222 Pa | Deer/km2 Number of plots SE 90% CI° required 1.9 32 160 1.1 14 85 0.6 31 190 0.9 49 375 0.004 30 190 0.03 14 100 0.1 is 125 0.1 13 120 0.2 15 105 0.5 4] 790 0.3 16 200 0.9 at 165 0.5 30 440 0.3 18 175 “A: systematic sampling of all forested areas in the zone. B: systematic sampling with survey limited to wintering areas > 2.5 km?. C: stratified sampling with survey of A and B strata. *Expressed as % of mean estimate. 2002 POTVIN, BRETON, AND RIVEST: DOUBLE-COUNT AERIAL SURVEY 493 TABLE 3. Deer density estimated by the double-count technique for eight aerial surveys replicated two to five times in Québec. Sighting probability Delay® of deer groups Deer/km? Survey Replicate (days) Front Rear m plots = 90% confidence limits Anticosti - Simone1988 1 0.48 0.63 20 15.6 11.5-19.7 2 2 0.49 0.63 20 13.2 10.1-16.3 3 4 0.55 0.60 20 12.3 8.2-16.4 4 7 0.70 0.78 20 11.6 9.9-13.3 Anticosti - Salmon 1988 1 0.65 0.59 23 99 7.8-12.0 2 2 0.53 0.67 25 8.5 6.3-10.7 3 F 0.73 0.58 ZS iis 8.4-14.2 Anticosti - Simone 1989 1 0.61 0.49 40 8.1 AB 6.1-10.1 2 7 0.47 0.36 40 11.7A 8.7-14.7 3 9 0.38 0.43 40 11.5B 8.3-14.7 Watopeka 1990 1 0.62 0.65 33 ay 2.8-6.6 2 6 0.75 0.88 28 3951) 1.9-5.9 3 31 0.39 0.56 28 2.6 CDEF 1.14.1 4 85 0.64 0.56 28 44E 2.2-6.6 5 86 0.43 0.55 28 4.1F 0.9-7.3 Armstrong 1991 1 0.67 0.56 35 11.6 8.7-14.5 2 64 0.50 0.60 38 10.9 8.0-13.8 St. Gédéon 1991 1 0.57 0.55 26 8.6 6.6—10.6 z 52) 0.24 0.29 pe 6.8 2.0-11.6 Ss 74 0.59 0.32 24 6.9 4.5-9.3 Island Brook 1991 1 0.85 0.75 SP 12.4 9.9-14.9 2 66 0.65 0.51 39 9.8 7.1-12.5 Témiscouata 1995 1 0.71 0.59 34 9.6G 6.9-12.3 2 7 0.29 0.12 33 6.6 G 2.2-11.0 “Number of days after the first replicate. >Within each survey, pairs of means having the same letter have a difference > 30% (biologically different). are used for deer than for Moose because deer occur at higher densities. For Moose, the plot size is 6 X 10 km (Créte et al. 1986). Therefore, total cost for a typical zone is about $43 000 for deer, as compared to some $150 000 for Moose (recalculated from Courtois et al. [1996] on a basis of $1000/hr for helicopter). Repeatability is important as conditions may change between surveys. Although no differences were detected among eight replicated surveys when comparing CI, this procedure had low power because of small sample size and high SE (Steidl et al. 1997). Nevertheless, only three of 24 replicates overall had a difference in deer density that we considered biologically significant (> 30% within each survey) with other replicates. The delay between replicates cannot explain such differences because no more than a month separated the replicates involved within these surveys. On Anticosti in 1989, the first replicate of Simone block provided a lower density than the fol- lowing two replicates. Because of thick and dense snow, deer were difficult to detect on that survey, and the observers possibly improved their skill on the sec- ond and third replicates under such conditions. In the Témiscouata wintering area, sighting probabilities of both observers were unusually poor (0.12-0.29) on the second replicate, when crusted snow was present and deer were immobile. This resulted in a low density estimate and poor precision (90% CI = 66%). Low sighting probability of the front observer (0.39) might also explain the results of the third replicate in the Watopeka wintering area, the only result that was dif- ferent in that survey. The problem in two of those three surveys is related to a low sighting probability (< 0.40). The sighting probability therefore appears to be a good measure of the quality of a double-count survey. Simulations have shown the technique to be robust to unequal capture probability (Magnuson et al. 1978), unless the sighting probability is < 0.45 (Caughley and Grice 1982). Graham and Bell (1989) suggested applying double surveys only to highly visi- ble (= 0.50) populations. Based on our replicated sur- veys, we suggest that results from the double-count technique are valid if sighting probability exceeds 0.45, but might underestimate deer densities when the sighting probability is < 0.40. As compared to the pellet-group count technique, double-count aerial surveys gave similar estimates in five of nine surveys of wintering areas. The largest difference stems from two surveys of the Poheéne- gamook wintering area, where the aerial estimate is twice the pellet-group estimate. Pohénégamook has 494 THE CANADIAN FIELD-NATURALIST S | Vol. 116 TABLE 4. Deer population estimated by the double-count aerial survey and by the pellet-group count techniques for nine deer wintering area surveys in Québec. Double-count aerial survey Survey m plots Population Armstrong 1989 25 2500 Watopeka 1990 89 860 Lac David 1990 60 1060 La Macaza 1990 62 2230 Armstrong 1991 35 1500 St. Gédéon 1991 26 1280 Island Brook 1991 32 1090 Pohénégamook 1994 61 480 Pohénégamook 1996 39 520 ‘Using a defecation rate of 26 pellet-groups/day. Pellet-group count 90% confidence limits population* 1780-3230 1770° 630-1090 680° 790-1320 1630° 1670-2790 2530° 1120-1870 1770° 970-1590 1040 870-1310 1190 430-530 3004 440-600 2504 ’Population estimate for winter 1990. No pellet-group count surveys were done in 1989 and 1991 but the estimates for 1988, 1990, and 1992 indicate a stable population. ‘Pellet-group count surveys. were done in 1989. ‘Dumont et al. (1998); population estimates by the pellet-group count were modified using a defecation rate of 26 pellet groups/day. the longest winters (>120 days) among our study areas and poor quality habitat (Dumont et al. 1998). Under such conditions and with reduced food intake, we suggest that the defecation rate might be lower than 26 pellet groups per day. In that same area, Potvin et al. (1981) conducted pellet-group counts and dead deer surveys from 1972 to 1979. Using the annual population estimate from the pellet count (defecation rate = 13 pellet groups per day) and the tallied number of dead deer, they evaluated winter mortality rates of up to 40%. With the same data, a defecation rate of 26 pellet groups per day would have lowered by half the population estimate and therefore doubled the mortality rate, which appears unrealistic. Therefore, a lower defecation rate certain- ly prevailed. Overall, we consider that the double- count aerial survey technique generally provides sim- ilar deer density estimates to pellet-group counts. Although the costs of both techniques are rather simi- lar, we suggest that aerial surveys provide more reli- able estimates because pellet-count evaluations require two arbitrary variables, defecation rate and deer concentration period inside the yard. Pellet-count estimates do not provide a valid stan- dard against which to measure accuracy of the dou- ble-count aerial survey technique. Because of plot size, an evaluation of the accuracy would require a very large enclosure (> 5 km?) with a known popula- tion, something rarely available for logistic and bud- get reasons. Using marked deer with radio-collars to measure sighting probability is neither feasible with double counts as the observer would not always be able to verify if the animal seen had a collar or not. Because deer densities are high, it is not possible to find the animal when flying back, which is the usual procedure for Moose. There are possible bias in the double-count tech- nique. First, the two observers aboard the aircraft have almost the same vantage point, so that the sighting probabilities may be overestimated and the true deer abundance underestimated. Furthermore, some concealed animals (e.g., under dense vegeta- tion) may have a null sighting probability (Marsh and Sinclair 1989), a factor also adding to negative bias. Overall, we consider the gradient of densities obtained in Québec by the double-count technique to be realistic. A general decrease in densities from southern to northern or eastern zones corresponds to observed winter severity and habitat quality. Anticosti Island is the only exception on that scale. Despite a boreal forest habitat, Anticosti has no predators and a low harvest rate (<10%), which explains the unusual high density at that latitude. Management Implications The double-count technique is reliable to survey White-Tailed Deer over medium and large areas at a reasonable cost, with great potential utility for state and provincial deer management programs. At north- ern latitudes, the best period is early winter when there is a uniform snow layer on the ground (2 10 cm). Later in winter, deer congregate in dense conif- erous cover, becoming less active and more difficult to see. Summer surveys can also be conducted where snow is absent, if there is a good contrast between the deer colour and the ground, and if deciduous trees and shrubs are not too dense. New Brunswick has already successfully used the technique in winter in parts of the province (Ballard et al. 1999; Gerry Redmond personal communication). Although it is feasible to manage deer populations by using only indirect data (e.g., harvest) and mod- els, these techniques are not quickly sensitive to drastic change. White-Tailed Deer are becoming a nuisance animal across North America (Warren 1997). Non-hunters and conservationists might more- 2002 POTVIN, BRETON, AND RIVEST: DOUBLE-COUNT AERIAL SURVEY 495 easily accept deer population reductions when deer numbers are obtained by precise aerial surveys rather than through pellet-group counts or harvest based computer models. Hunters also might be more will- ing to increase or decrease harvests if reliable deer density figures are provided. Acknowledgments We wish to thank all regional personnel of Société de la faune et des parcs du Québec that participated in planning and conducting surveys. Michel Créte and four anonymous reviewers provided helpful comments on a previous version of the manuscript. Literature Cited Ballard, W. B., H. A. Whitlaw, S. J. Young, R. A. Jenkins, and G. J. Forbes. 1999. Predation and survival of white-tailed deer in northcentral New Brunswick. Journal of Wildlife Management 63: 574-579. Bartmann, R. M., L. H. Carpenter, R. A. Garrott, and D. C. Bowden. 1986. Accuracy of helicopter counts of mule deer in pinyon-juniper woodland. Wildlife Society Bulletin 14: 356-363. Bartmann, R. M., G. C. White, L. H. Carpenter, and R. A. Garrott. 1987. Aerial mark-recapture estimates of confined mule deer in pinyon-juniper woodland. Journal of Wildlife Management 51: 41-46. Beasom, S. L. 1979. Precision in helicopter censusing of white-tailed deer. Journal of Wildlife Management 43: 777-780. Beasom, S. L., J. C. Hood, and J. R. Cain. 1981. The effect of strip width on helicopter censusing of deer. Journal of Range Management 34: 36-37. Beringer, J., L. P. Hansen, and O. Sexton. 1998. Detection rates of white-tailed deer with a helicopter over snow. Wildlife Society Bulletin 26: 24-28. Boer, A. H., G. Redmond, and T. J. Pettigrew. 1989. Loran-C: a navigation aid for aerial surveys. Journal of Wildlife Management 53: 228-230. Breton, L., and F. Potvin. 1997. Normes d'inventaire aérien des populations de cerf de Virginie. Québec Ministére de l'Environnement et de la Faune, Report 3712-97-09. 44 pages. Caughley, G., and D. Grice. 1982. A correction factor for counting emus from the air, and its application to counts in western Australia. Australia Wildlife Research 9: 253-259. Cochran, W. G. 1977. Sampling techniques. John Wiley & Sons, New York. 413 pages. Courtois, R., Y. Leblanc, J. Maltais, and H. Crépeau. 1994. Québec moose aerial surveys: methods to esti- mate population characteristics and improved sampling strategies. Alces 30: 159-171. Courtois, R., F. Potvin, S. Couturier, and A. Gingras. 1996. Révision des programmes d'inventaires aériens des grands cervidés. Québec Ministére de l'Environ- nement et de la Faune, Report 96-3425-10. 49 pages. Couturier, S., R. Courtois, H. Crépeau, L. P. Rivest, and S. Luttich. 1996. Calving photocensus of the Riviére George caribou herd and comparison with an indepen- dent census. 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C., S. D. Dubois, D. J. Reed, and S. J. Harbo. 1986. Estimating moose population parameters from aerial surveys. University of Alaska, Biological Paper 22. 108 pages. Gasaway, W. C., and S. D. Dubois. 1987. Estimating moose population parameters. Swedish Wildlife Research, Supplement 1: 603-617. Gill, R. M. A., M. L. Thomas, and D. Stocker. 1997. The use of portable thermal imaging for estimating deer population density in forest habitats. Journal of Applied Ecology 34: 1273-1286. Graham, A., and R. Bell. 1989. Investigating observer bias in aerial survey by simultaneous double-counts. Journal of Wildlife Management 53: 1009-1016. Havens, K. J., and E. J. Sharp. 1998. Using thermal imagery in the aerial survey of animals. Wildlife Society Bulletin 26: 17-23. Kilpatrick, H., and M. Ellingwood. 1993. New method used in 1993 aerial deer survey. Connecticut Wildlife 13(4): 3. Kufeld, R. C., J. H. Olterman, and D. C. Bowden. 1980. A helicopter quadrat census for mule deer on Uncompahgre Plateau, Colorado. Journal of Wildlife Management 44: 632-639. Lamontagne, G., and F. Potvin. 1994. Plan de gestion du cerf de Virginie au Québec 1995-1999 — L'espéce, son habitat et sa gestion. Québec Ministére de I'Environ- nement et de la Faune, Report 94-2501-11. 114 pages. Lancia, R. A., J. D. Nichols, and K. H. Pollock. 1994. Estimating the number of animals in wildlife populations. Pages 215-253 in Research and management techniques for wildlife and habitats. Edited by T. A. Bookhout. 5th edition. The Wildlife Society, Bethesda, Maryland. Leptich, D. J., D. G. Beck, and D. E. Beaver. 1994. Aircraft based Loran-C and GPS accuracy for wildlife research on inland study sites. Wildlife Society Bulletin 22: 561-565. Magnusson, W. E., G. J. Caughley, and G. C. Grigg. 1978. A double-survey estimate of population size from incomplete counts. Journal of Wildlife Management 42: 174-176. Marsh, H., and D. F. Sinclair. 1989. Correcting for visi- bility bias in strip transect aerial surveys of aquatic fauna. Journal of Wildlife Management 53: 1017-1024. Neff, D. J. 1968. The pellet-group count technique for big game trend, census, and distribution: a review. Journal of Wildlife Management 32: 597-614. 496 Pollock, K. H., and W. L. Kendall. 1987. Visibility bias in aerial surveys: a review of estimation procedures. Journal of Wildlife Management 51: 502-510. Potvin, F. 1995. L'inventaire du brout: revue des méth- odes et description des deux techniques. Québec Ministére de l'Environnement et de la Faune, Report 95-2545-02. 70 pages. Potvin, F., J. Huot, and F. Duchesneau. 1981. Deer mor- tality in the Pohénégamook wintering area, Québec. Canadian Field-Naturalist 95: 80-84. Potvin, F., L. Breton, L. P. Rivest, and A. Gingras. 1992. Application of a double-count aerial survey tech- nique for white-tailed deer, Odocoileus virginianus, on Anticosti island, Québec. Canadian Field-Naturalist 106: 435-442. Rivest, L. P., F. Potvin, H. Crépeau, and G. Daigle. 1995. Statistical methods for aerial surveys using the double-count technique to correct visibility bias. Biometrics 51: 461-470. ° Rogers, L. L. 1987. Seasonal changes in defecation rates of free-ranging white-tailed deer. Journal of Wildlife Management 51: 330-333. Sawyer, T. G., R. L. Marchinton, and W. MacLentz. 1990. Defecation rates of female white-tailed deer in Georgia. Wildlife Society Bulletin 18: 16-18. THE CANADIAN FIELD-NATURALIST : ? Vol. 116 Seber, G. A. F. 1982. The estimation of animal abun- dance and related parameters. 2nd edition. MacMillan Publishing Company, Inc., New York. 653 pages. Seber, G. A. F. 1992. A review of estimating animal abundance II. International Statistical Review 60: 129-166. Steidl, R. J., J. P. Hayes, and E. Schauber. 1997. Statistical power analysis in wildlife research. Journal of Wildlife Management 61: 270-279. Stoll, R. J.. M. W. McClain, J. C. Clem, and T. Plageman. 1991. Accuracy of helicopter counts of white-tailed deer in western Ohio farmland. Wildlife Society Bulletin 19: 309-314. Tzilkowski, W. M., and G. L. Storm. 1993. Detecting change using repeated measure analysis: white-tailed deer abundance at Gettysburg National Military Park. Wildlife Society Bulletin 21: 411-414. Warren, R. J. 1997. The challenge of deer abundance in the 21st century. Wildlife Society Bulletin 25: 213-214. White, G. C., R. M. Bartmann, L. H. Carpenter, and R. A. Garrot. 1989. Evaluation of aerial line transect for estimating mule deer densities. Journal of Wildlife Management 53: 625-635. Received 10 November 2001 Accepted 9 November 2002 Book Reviews ZOOLOGY Survivors in Armor: Turtles, Tortoises, and Terrapins Orenstein, Ronald. 2001. Key Porter Books, Toronto. xi + 308 pp. Illus., $45. With their armour of carapace and plastron, turtles are one of the most distinctive groups of organisms in the world. Yet few naturalists seem to know very much about turtles. Ronald Orenstein has set out to correct this oversight in this wonderfully fascinating and beautiful book. In ten chapters Orenstein covers all the major top- ics of turtle biology and conservation. The book begins with a general introduction to the diversity of turtles and their ecological complexity. Next, a thor- ough analysis of turtle evolution clearly charts out the competing theories of turtle ancestry. Two chap- ters provide a summary of each of the families of turtles, allowing the reader to see patterns within groups of turtles. The next few chapters cover vari- ous aspects of turtle physiology, ecology, and repro- duction. The book concludes with two chapters on threats to turtles: one on sea turtles and one on all other species. Orenstein freely admits he is not a herpetologist, but he had three expert consulting editors: Jean Mortimer, Chair of the Hawksbill Task Force; Peter Pritchard, Director of the Chelonian Research Institute; and George Zug, Curator of Herpetology at the National Museum of History in Washington DC. This combination of talent has produced a book that is highly detailed in information, yet clearly written and aimed at a general audience. The book is also lavishly illustrated with colour photographs. The quality of the photos ranges from good to exception- al and they feature many of the species mentioned in the text. Although many of the species featured are from North America there are also photos of species from Asia, Australia, and South America. The book is filled with wonderful little gems of detail that illustrate the incredible diversity of life- history strategies within this small group of organ- isms. For example, the Tent Tortoise (Psammobates tentorius) of South Africa has evolved “gutters” along the edge of the carapace. During light rains or mists, water collects on the carapace, runs into the gutters, and, by tilting its carapace forward, water runs forward and the tortoise can drink. And the Northern Long-necked Turtle (Chelodina rugosa) of Australia has the distinction of being the only known reptile which lays its eggs under water. The eggs are laid underground in temporary ponds, don’t start developing until after the water dries up, and don’t hatch until the start of the next rainy season. Turtles face horrific threats to their existence and Orenstein does a thorough job of describing their bleak future. Great emphasis is placed on the current crisis in the trade in Asian turtles for food. Over nine million live turtles were imported into Hong Kong in 1998 alone. Approximately half of the 89 species of turtles native to Asia are considered to be endan- gered or threatened. North American species are not much better off with populations threatened by habit loss, traffic mortality and collecting for the pet trade. Sea turtles have been overharvested for centuries and Orenstein illustrates this effectively with the tale of Bermuda. Green Sea Turtles were so depleted around Bermuda that the Bermuda Assembly passed laws to protect them as early as 1620. Despite such attempts it is now estimated that Green Sea Turtle populations in the Caribbean have decline by at least 99% since the time of Columbus. Orenstein also dis- cusses other threats such as fibropapillomatosis (a mysterious disease that causes fibrous tumors), long- line fishing vessels, and shrimp trawlers. He also explores the complex political waters of the WTO (World Trade Organization) and CITES (Convention on International Trade in Endangered Species) that sea turtles have had to navigate. Why should we care about the fate of these rep- tiles? As the author states in his preface, turtles “matter because it would be shameful if their long tread through 200 million years of evolutionary his- tory should end through our negligence, our greed, and our failure to act” (page xi). This book is a cele- bration of the wonder of turtles and a plea for their continued survival. I hope it succeeds. DAVID SEBURN Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford Mills, Ontario KOG 1S0 Canada 497 498 THE CANADIAN FIELD-NATURALIST . a ea aman Ge Ss) oa ow wel ue he ine) Vol. 116 Amphibians and Reptiles of Pennsylvania and the Northeast By Arthur C. Hulse, C. J. McCoy, and Ellen J. Censky. 2001. Comstock Publishing Associates, a division of Cornell University Press, Ithaca. xi + 419 pp. Illus., U.S. $39.95. In theory, this substantial volume should be the definitive guide to the 83 species of amphibians and reptiles found in the northeastern United States. It should also be a valuable resource for eastern Canadian naturalists as most of the amphibians and reptiles found in Ontario or to the east are covered in this book. Unfortunately this book falls far short of the standard expected from a university press. The structure of the book is straightforward but thorough. The book opens with a brief introduction on the geography and climate of the area. Next a set of keys is provided for both larval amphibians and adult amphibian and reptiles. The book also includes 133 colour photos of the species covered. The photos are small (four or more per page) but there is more than one photo for many species. These additional photos are often of colour variations, eggs, or larvae. The bulk of the text is devoted to the species accounts. Accounts vary from approximately 2—4 pages in length and include a distribution map (locality dots for Pennsylvania, but just shaded for the rest of the north- eastern range). Each account follows the same stan- dard format with the following headings: description, confusing species, habitat and habits, reproduction, remarks, and distribution. The book concludes with an appendix, glossary, and literature cited. The appendix is a compilation of measurements on males and females of most species from Pennsylvania. For example, for salamanders they provide the mean (+ standard error) snout-vent length and total length of males and females of each species and note whether the differences are statistically significant. The outline for this book is thus detailed and thor- ough. Unfortunately, the execution leaves much to be desired. The worst of the mistakes are in the keys. Although profusely illustrated the labels sometimes are in error. For example, the illustrations of ven- omous vs. non-venomous snakes are switched. The The Sibley Guide to Bird Life and Behavior Illustrated by David Allen Sibley, edited by Chris Elphick, John B. Dunning, Jr., and David Allen Sibley. 2001. Alfred A. Knopf Inc., New York 608 pp. illus. U.S. $45 This new Sibley guide is a useful contribution to the rather limited number of volumes that seek to take the birder beyond identification to the complex- ities of “how birds live and what they do.” It features Sibley’s excellent artwork throughout, and he is one of the editors, but the actual text is the work of over 40 individual authors. key to adult salamanders has a critical typo making it impossible to get past step eight and into the plethodon salamanders. Including a key to tadpoles is wonderful, but for some reason the authors omit- ted three species (Pine Barrens Treefrog, Mink Frog, and Carpenter Frog) with no explanation. The species accounts are generally fairly good as far as they go. More information could be provided on these species, but space limitations obviously had to be balanced with the amount of detail. An extra page on each species would have added almost 100 pages to the final book. There are a few factual errors. The authors state it is unknown whether Pickerel Frogs hibernate on land or in water, yet this species is known to generally hibernate under water. There are also some peculiar omissions. The Wood Turtle account fails to mention worm-stomping, surely one of the most intriguing feeding behaviours in any tur- tle, despite the fact that the discovery was made in Pennsylvania. The most surprising gap in this book is any sub- stantial treatment of amphibian decline, or conserva- tion in general. There is no single section on the growing threats to amphibians and reptiles, only occasional references in the remarks section of the species accounts. Even these occurrences are some- times bewildering. For example, the authors write “Although the wood turtle is not considered to be in any immediate danger in Pennsylvania, the potential for extirpation in the state does exist” (p. 214). Apparently potential extirpation from the entire state is not “immediate” danger. In summary, while this book had the potential to be an excellent resource on amphibians and reptiles it ends up being a flawed but moderately useful book. Wait for the second edition and hope the authors pay more careful attention to detail. DAVID SEBURN Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford Mills, Ontario KOG 1S0 Canada The book is divided into two broad sections. Part I, some 106 pages long, consists of five chapters on the “World of Birds”, and covers subjects ranging from Flight, Form, and Function, to Behavior, Populations, and Conservation. Each chapter is well organized with the various topics discussed under a ‘series of sub-headings, and taken together, they pro- vide a comprehensive survey of birdlife in a concise and readable format. Part II, and the greater bulk of the book at some 2002 BOOK REVIEWS 499 440 pages, is a survey of the bird families of North America. Each family is covered in turn, with the treatments ranging from two to 20 pages, depending on the number of species in the family. The approach mirrors Part I, with subject headings on Taxonomy, Food and Feeding, Breeding, Movements, and Conservation, and additional head- ings for the larger chapters. Each family account also includes a box giving a summary of “Worldwide Family Features”’. Because each section has its own authors there is some unevenness in style, and some parts are heavier than others, but in general the editors have done an excellent job of producing a book that is free of jar- gon and easily read. I did find the odd over-general- ization or misstatement, but these were few and in general the book was pleasantly error-free. There is a comprehensive index that seems to work well, and which allows the reader to follow up on any refer- ences to particular species in the text. The illustra- tions are well planned, and the book concludes with a glossary and a checklist of species. Persons familiar with the introductory sections of the landmark Handbook of the Birds of the World series will see an immediate parallel in approach, but I know of no books on this continent’s birdlife that provide this kind of broad overview. Most of the other books of this genre approach the topic from the level of the individual species — for example, The Birder’s Handbook, by Ehrlich et al., which has been an invaluable source of concise natural history infor- mation for over a decade. The question is whether a book on families provides a new and valuable approach, or whether it is simply too general to give more than a superficial overview. Certainly much of the treatment is necessarily quite general. But — because we naturally tend to concentrate more on the individual species than on the families they belong to — we often can overlook the broader picture. It is exactly this broader per- spective that this guide now provides. This is a par- ticularly useful approach for the average birder, who has often developed his or her knowledge and skills working with the traditional field guides, and who may have only a fragmentary understanding of the broader patterns of relationships within and between bird families. Such information can often be helpful with finding the birds in the first place and under- standing them better, but it really comes into its own if the individual becomes involved in some other aspect of bird life, such as atlassing. Its utility is certainly not confined to the newcom- er. Even the experienced birder will probably find new information here — you may think you know all this stuff, but you may be surprised at how much you don’t know, or have forgotten! For the veteran it forms a fast reference and a very readable summary of the current information on bird life. All in all, this is a very worthwhile contribution to the North American bird literature. It brings together an up-to-date body of information that is not readily available in one place elsewhere, and presents it in an attractive, readily understandable form. If you want to expand your horizons beyond simply identi- fying your birds, this is a book to buy. CLIVE E. GOODWIN 1 Queen Street, Suite 401, Cobourg, Ontario K9A 1M8 Canada A Birder’s Guide to Metropolitan Areas of North America Edited by Paul Lehman. 2002. American Birding Association Inc., Colorado Springs, Colorado. iv + 508 pp., illus. $US. $26.95. I travel a fair amount for business and so I look for birding opportunities wherever I go. For some years I thought I should write a guide for people who, like me, spend a day or two in a city and then move on. Now Paul Lehman has produced such a book. He has collected a group of local experts or groups to write a chapter on 33 major North American cities (including seven Canadian cities). This book is written in the same pattern as the other ABA/Lane guides. For each city there is a description of any good birding sites within the city and surrounding area. This account includes the most interesting species present with due reference to seasonal changes. I thought the descriptions of the birds you are likely to find were realistic. Too often I read an account of a birding “paradise” only to find that it includes twenty years worth of rarities as if they were regular visitors. This is not to say the mega-finds are not included, but they are kept in context. Clear maps and detailed instructions, including parking spots, are provided to assist the visitor in finding their way around. In addition to the information on birds there are also useful cli- mate and travel tips, plus material on driving times and safety. The content of the book was provided by some of the most experienced and knowledgeable people in each area and reviewed by an equally impressive group of experts. Although this is a very useful guide it is not quite what I had in mind. I was thinking of a book that would describe areas you could walk to from a downtown hotel (or at least reach with public transit or a short cab ride). This book covers not only the inner parks and shorelines but has a lot of informa- tion on areas that are one to two hours drive away. 500 For example, the Detroit section includes Port Huron and Calgary includes Peter Lougheed Park; both about an hour and a half drive. This is not what I think of as a “metropolitan” area, so for many of the areas mentioned you will need a car. Although the editor states this book is intended for visitors to the downtown core and those who can wander farther afield, the bias is for the mobile birder. There are many cities missing, Victoria, British Columbia and St Johns, Newfoundland are two that come immediately to mind, as they are both places where you can easily walk to good birding areas from downtown. The biggest gap though is in the prairie and central plains region and, less surprising, there is nothing north of Calgary, Alberta. Book size is always an issue when creating a portable guide. This 33 city volume is already roughly equivalent to a National Geographic Field Guide to Birds of North America. It will, therefore, easily fit in your travel case. Personally, I am more likely to photocopy a The World of the Hummingbird By Robert Burton. 2002. Firefly Books Ltd., Willowdale, Ontario, Canada. 158 pp., illus. $40. The philosophy of the wise prairie naturalist Isabel Priestly was that articles (or books) should be written as if two naturalists were exchanging their experiences in an informal way, yet were scientifi- cally correct. Robert Burton is definitely that kind of naturalist and author. He is a practiced and skilled writer who is able to tell a vivid tale, underpinned by the best of scientific knowledge. In this book, his storytelling ability not only applies to the joys of seeing this extraordinary group of birds, but to scien- tific explanations of complex phenomena. For exam- ple, the author evidently has a keen interest in the physics of flight. His explanations of hummingbird flight are clear, straightforward and accurate and very understandable. He covers the fundamentals of colour in hummingbird feathers (or any other bird for that matter) plainly and perfectly, while retaining the magic the colours evoke. There is much about hummingbirds that is well studied. These birds attract researchers not simply for their colourful plumage, but by their unusual behaviour and their visibility. The author has drawn from these studies to give us a glimpse into the life of these tiny birds. In addition to a general introduc- tion and the aforementioned section on flight, the author explains the all-important relationship of these birds with flowers. The influence that flowers have on the size and shape of each species and the adaptations of the flowers to accommodate their principal agents for pollination make enticing read- ing. The chapter on the social behaviour I found fas- THE CANADIAN FIELD-NATURALIST 1 ——— i ame Vol. 116 - few relevant pages to take with me. These I couid write on, drop in the mud, and eventually discard without disturbing my original text. Like any book of this type, it starts going out of date as soon as it is published. For example Shepard Slough near Calgary has been drained and the Crested Mynas on Wylie Street, Vancouver are down to a single pair (currently nesting). To combat this problem there are resources such as web sites, RBAs, and phone numbers to bring you up-to-date. The concept of this book is good and in its current form will be of great value to the travelling birder. I hope that the process is continued and a second vol- ume or edition will fill in some of the blanks. Now how do I justify a trip to Los Angeles? Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada cinating and, for me, threw new light on the role played by the hummingbird’s belligerent behaviour . Despite their minuscule size hummingbirds make some prodigious flights. As the author explains these not only follow the seasons but track the food supply as well. The nesting biology is both surprising and engag- ing. Their diminutive nests are remarkable construc- tions that are often difficult to spot (I once spent 40 minutes pointing out a nest to a large group, and many did not see it until the female landed on it). The strategies the birds use to ensure nesting success are equally remarkable. The author finishes with the relationship of hum- mingbirds to humans. I was astounded to read that in the 17" century hummingbird skins sold for £8.00 (about $2000 in today’s money) and ONE dealer imported 400 000 skins in ONE year. It is amazing there are still hummingbirds left. Happily times have changed and hummingbirds get some measure of protection, and in some places considerable help. The book is illustrated by some frame-filling, crisply focussed photographs. Given the bird’s inher- ent twinkling brilliancy and given that they are typi- cally feeding on a sumptuous flower these pho- tographs are just delightful. This is one of the best-written books I have read. It would be a wonderful gift, even if you give it to yourself. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada 2002 BOOK REVIEWS 501 The Complete Encyclopedia — Antarctica and the Arctic By D. McGonigal and L. Woodworth. Firefly Books Inc., Willowdale, Ontario. 608 pp. Illus. Cloth $59. This is a wonderful book that captures the Antarctic experience remarkably well. It is a large- sized volume with roughly 50% of the space taken by photographs, much like a coffee table book. It also has a substantial amount of explanatory text that makes it far more than a simple picture book. While I might question the term “complete encyclopedia,” the book does have a great deal of substance. For reasons that will emerge below, I think it is best to emphasize the Antarctic content first. The book is divided into six parts. The first three parts cover the natural environment, including wildlife, climate, geology, and geography. The sec- ond half of the book deals with the various compo- nents of human interaction with the Polar Regions. The explanations begin with a description of the current and evolutionary geology of the two poles, followed by an explanation of the climate. In partic- ular, the various stages of sea ice formation and the birth of icebergs are described in some detail. Human influence on climate change (including the author’s polite use of “burp” for the emission of methane by our cattle) together with the concerns over the ozone hole are explored in precise tones that even an American president will understand. The geographical section describes the key fea- tures of the entire Antarctic region. Something of the history and, hence, the names of various areas along with some geological background are given. Also included, justifiably in my opinion, are the sub- Antarctic islands. Though they lie beyond the Antarctic proper, they have played a major part in the region’s natural and human history. More than half the book is devoted to Antarctic Wildlife and Exploration. As well as giving a gener- al introduction to Antarctic ecology and the role played by krill and other marine life, the authors give detailed accounts of most of the prominent wildlife. Seals, cetaceans, and birds are treated in the same manner as a reference text such as Birds of Canada’. Penguins, as the real symbol of this continent, rate a chapter of their own. Range maps are included where appropriate. The second major section is on Antarctic explo- ration. This is a thorough and well-written review of this exciting phase of human endevour. The early whalers are treated fairly for their exploration and contribution to knowledge without being judgmental on morals that existed 150 years ago. All the well- known names appear under the heading “Heroic Age”; Shackleton, Scott, Amundsen, etc. But other lesser lights like Nobu Shirase and William Filchner get an appropriate and respectable mention. As well as current photographs this section contains repro- ductions of original plates taken by the expedition photographers. There are some beautiful renditions of Frank Hurley’s classical photos of the ill-fated Shackleton expedition of 1914. The text is profusely illustrated with photographs that range from good to stunning, with most being at the high end of the scale. The photographs of nature are simply wonderful and they invade the non- wildlife sections of the text. Here their sole function seems to be decorative. They do not appear to relate to the text, but why should we complain about such lovely images? They are just delightful. The illustra- tions include a number of beautiful, clear, and accu- rate maps. I was impressed to see South Georgia correctly positioned. Many years ago someone pro- duced a map where South Georgia was 1° in error and this mistake has been reproduced time and again. Now let us look at the Arctic portion of this ency- clopedia. While the material devoted specifically to the Antarctic takes up about 70 percent of the pages, the Arctic gets about 10 percent. This is an enormous disparity and it shows. In the wildlife section only seven species of birds are depicted and a number of famed Arctic residents get little or no mention. What has happened to the geese, eider, shorebirds, falcons, and pipits, not to mention the most famous species of them all, Ross’s Gull? The mammal section is also very short, despite being padded by some “southern- ers” like Wapiti (now virtually confined to the south- ern Rockies). Oddly there is no photograph of Caribou, except a picture of Reindeer taken in South Georgia. The Moose shown is found a little closer to the Arctic (southern Hudson’s Bay and it has been pushing northwards in Labrador), but I hardly think of it with Polar Bears and Grey Wolves. The marine mammals fare a little better, but hardly get equivalent treatment to their southern counterparts. The section on Arctic Exploration is similarly short. It does mention the key figures such as Frobisher, Parry, Franklin, and Nansen, but there are a number of people missing. Three in particular seemed notably absent. Touching my national pride a little I missed seeing mention of Bob Bartlett of Newfoundland, Canada. Over 40 voyages and more than 200 000 miles sailing in Arctic waters should get at least an honourable mention. Captain Henry Larsen’s two historical traverses of the North-west Passage in the St. Roch — west to east in 1942 and east to west in 1944 — is another notable Canadian achievement neglected by the authors. A further important fumble is leaving out Ralph Plaisted of Minnesota. Plaisted and three others were the first men (1968) to make an overland trip to the North Pole that has indisputable verification (by two sepa- rate US Airforce navigators using gyro-compasses). Peary and Cook’s claims are far too unrealistic to be acceptable. Among the other notables not mentioned are Bylot, Sverdrup, and Papinin. i — a Ti) Te wR PRVAN ° 502 There are a few minor errors. “Ruffled” Grouse — part of an insert on cyclical food chains — is clearly a simple mistake. Making a point of calling a Yellow- billed Pintail carnivorous is a bit misleading. There are many carnivorous ducks (mergansers, goldeneye etc.); this pintail has the bizarre habit for a duck of scavenging corpses. The authors start their discussions on cormorants and then switch (correctly) to the term “shags” but without an explanation of the difference. The text on skuas does not cover the complexity and disagreement of this difficult group. Some of the pho- tos do not have precise identifications. This tome concludes with a section on living in the Polar Regions (mostly the Antarctic though) and some of the activities that are taking place (research, aviation, shipping, fishing, hunting, and tourism). The Complete Guide to the Birds of Europe By L. Svensson and P. Grant (Translation by D. Christie, originally published in Swedish in 1999). 2002. Princeton University Press, Princeton and Oxford. 399 pp., illus. £25. In 1977 the Royal Society for the Protection of Birds (RSPB) published a booklet by M. Everett & P. Hayman called What’s That Bird'. This publica- tion consisted mostly of illustration with little text. It showed birds in classical field guide format (mostly side views) and added numerous vignettes of birds flying away, in silhouette, front view and so on. From this multiplicity of images the reader got a much better idea of that quality birders call “jizz.” I long wondered why this novel idea was not imitated. Now it has been copied in both North America and with this book, Europe. In my opinion there are four development stages birders go through. The first is to identify birds by field marks, in the Peterson style. Then birders learn about habits and habitat. Next they work at identifying birds by song. Finally, the real enthusiast gains suffi- cient experience to use “jizz” — a combination of subtle features like relative body proportions, nature of flight, oddities of behaviour and a multitude of minute details that demand dedicated observation. We may have to invent a new term for this kind of book. It is too large (8.5’> X 12” X 1” or 22cm X 31cm X 2.8 cm) to be considered a field guide. Yet it is much smaller than Birds of the Western Palearctic? and similar reference works. How about calling it a reference guide? It is similar in style to a field guide, but because of its much larger format it contains about 3.5 times more illustrations than a typical guide does. For example, for most species there are roughly six depictions per species versus three in the National Geographic} and less than two in Peterson‘ This allows the authors to show the dif- THE CANADIAN FIELD-NATURALIST Vol. 116 - The final entry is a full text of the Antarctic treaty and Antarctic websites. So this magnificently illustrated book does a won- derful job on describing the Antarctic and gives a very brief introduction to the Arctic. As such it is a great resource for basic material on the Antarctic. While it is worth buying for the pictures alone, it does contain an “encyclopedic” amount of well- written information too. 'Birds of Canada. Revised Edition. By Earl Godfrey. 1986 National Museums of Canada Bulletin Number 203. Ottawa, Canada. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada ferent look a bird could have due to light or position or subspecies. In other words it gives a better sense of its “jizz.” The main illustrations are also around 1.5 times larger than other comparable books. Moreover there is about twice as much supporting text than the typical guide. The range maps are good, but at one inch square are rather small. The artwork, by Killian Mullarney and Dan Zetterstr6m, is impressive. I saw some of Mullarney’s work when he was still young and was struck by his skill. His partner Zetterstr6m has a similar format and quality so the book is visually smooth. Except, that is, for one discordant note. The illustrations of the North American warblers are by Larry McQueen. Although well done they are in a different style to the rest of the book. The two main artists have contributed many excellent illustrations of other North American birds (Gulls, thrushes, shorebirds, ducks etc.) so I was sur- prised at their reluctance to paint the warblers too. I thought the gull plates were very good, showing some of the best information on Common vs. Ring- billed gull and Herring vs. Ring-billed gull. There was no depiction of the pink flush on Franklin’s Gull. It is one of the few texts that show the juvenile plumage of Little Gull that can be confused with Sabine’s Gull — a mistake I have seen happen. Both male and female Rough-legged Hawks are shown. There is a plate dedicated to duck hybrids and a spe- cial section on the different diving techniques of scoters. In my copy, there are a few pages where some, but not all, birds are too pale, including some geese, larks and pipits. There are some species where the colour is too intense. These include Evening Grosbeak and some owls. The female Barrow’s Goldeneye beak, one of its key features, is far too dark. Virtually the entire bill is a rich golden yellow, 2002 BOOK REVIEWS 503 a mark I showed a visitor even in poor light this win- ter. The shapes of birds are particularly accurate. The only one I thought was too stocky was the should- be-sleek Black-browed Albatross. The Great Grey Owl shows a streaked breast with no horizontal bar- ring. This is the Strix nebulosa lapponica race that will have a pale appearance to North Americans. The back of the Hawk Owl has large pale patches which mark as the nominate race Surnia ulula ulula and this will also look a little odd to North Americans. A few of the extreme rarities and introduced species are not illustrated. The author’s choice of English names is a bit con- fusing. They start by using Red-throated Loon not Diver, but go to Great Northern — not Common — Loon. They add such words as (Common) in paren- theses in front of Raven, but also in front of (Black- billed) Magpie. Hoary Redpoll is called an Arctic and White—winged Crossbill is a Two-barred. The scientific names are, of course, consistent. Overall, this is an excellent book not only for Europeans but for North Americans too. Many birds from North America are covered and covered well. For example, there is good information on separating Grey-cheeked and Swainson’s thrush. As well, of course, there are numerous European species that occur here, such as Fieldfare, that get detailed treat- ment. It is a fine reference for all those species the two continents have in common. References What's That Bird.” M. Everett & P. Hayman. 1977. RSPB *Handbook of the Birds of Europe, the Middle East and North Africa — Birds of the Western Palearctic. S. Cramp. 1980 Oxford University Press 3Field Guide to the Birds of North America. National Geographic Society. +A Field Guide to the Birds of Britain and Europe. R. Peterson, G. Mountfort and P. Hollum. 1954. Collins. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada The Cuban Treefrog in Florida: Life History of a Successful Colonizing Species By Walter E. Meshaka Jr. 2001. University Press of Florida, Gainesville. xxiii + 191 pp. Illus., U.S. $69.95 The Cuban Treefrog (Osteopilus septentrionalis) is a hylid treefrog native to Cuba and the Bahamas. It appears to have been introduced into southern Florida early in the 20th century and by the mid- 1970s its distribution was fairly continuous through- out south-central Florida. This alien invader is not alone: to date there are over 30 exotic species of amphibians and reptiles established in Florida. The question is why are some species successful colonizers and others are not. Meshaka attempted to answer this question, with regards to the Cuban Treefrog, in his PhD dissertation, which focussed on Everglades National Park. He examined six general aspects of the Cuban Treefrog’s biology that pre- sumably affected its colonizing ability: reproduc- tion, activity patterns, habitat affinity, diet, preda- tors, and body size. He then examined each of these characteristics with ten ecological correlates associ- ated with successful colonization: high fecundity, short generation time, ability to function in a wide range of physical conditions, similar habitats in native and introduced ranges, coexistence with humans, broad diet, open niche space, superior com- petitive ability, predator-free space, and large body size compared with its closest relatives. The Cuban Treefrog met all but one of the ecological correlates. The niche space was not open, but this did not impede its ability to colonize. Cuban Treefrogs lay many eggs, can breed at almost any time of the year in southern Florida, flourish in disturbed environ- ments and can eat virtually anything, including many of their competitors. Meshaka has done a lot of worthwhile, detailed work. He has dissected hundreds of Cuban Treefrogs to examine their reproductive state across the sea- sons, counting ovarian eggs, and identifying prey from their guts. It nonetheless makes for a slim vol- ume that has been padded to reach even this length. For example, in one chapter he includes over 30 mediocre black-and-white photographs of Ever- glades National Park. Many of the photos provide before and after shots of the effects of Hurricane Andrew. This is worthwhile, but the author just doesn’t know when to stop. Likewise, Meshaka puts in every conceivable figure and table he can think of including one illustrating the accumulation curve for papers published on the Cuban Treefrog from 1750 to the present. The chapter on diet reaches a crescen- do with 24 figures and 29 tables. Detailed work that should definitely be in his dissertation, but one expects some sieving of material before publication. Of course that means there wouldn’t be enough material for a book, but there would be two or three strong papers for publication. And given the hefty price for this book that route would have likely reached a wider audience. DAVID SEBURN Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford Mills, Ontario KOG 1S0 Canada _— TO LR Ny OR 504 THE CANADIAN FIELD-NATURALIST Vol. 116 National Audubon Society Guide to Marine Mammals of the World By P. A. Folkens, R. R. Reeves, B. S. Stewart, P. J. Clapham, and J. J. Powell. 2002. Alfred A. Knopf Publisher, New York. 527 pp., illus. U.S.$26.95. This is another milestone publication and a won- derful and handy guide book which is full of details about the fascinating world of sea mammals. It was created by a high-caliber team of five North American authors with world-wide experience. Forget about “World-Birding”; instead, this book presents the solid foundation for “World-Whale- Watching”: the world’s sea mammals (“whales, dol- phins, porpoises, seals and sea lions, manatees, Marine and Sea Otters, and the Polar Bear’’) all in one solid book. The user will agree that this “field guide” turns easily into a fascinating read. A true highlight of this book are its 320 splendid drawings by Pieter Folkens; but the gorgeous 418 photographs, many from field research world-wide, deserve major recognition as well. Although most whale watchers will not see animals “underwater”, the stunning illustrations cover sea mammals in and out of the water, and show in detail the special fea- tures of sea mammals. A short chapter about Marine Mammal morpholo- gy explains all body features of a “sea mammal”. The authors introduce topics like “What is a Marine Mammal”, “Behaviour”, “Reproduction”, “Food and Foraging”, “Status and Conservation,” and “Watch- ing Marine Mammals” (e.g., Equipment, Etiquette, and Stranded Marine Mammals). As in other field guides, such topics are “kept on the surface’. I find the balance between information and usability in the field is well achieved. The 120 species accounts cover much ground. They present explanations of sea mammal names, number of teeth (no formula given), species descrip- tion, similar species, range and habitat, 123 distribu- tion maps (coordinate system included), behaviour, baseline data, and status and conservation. Species morphs are covered, so are subspecies (but their sci- entific names are not always given). The species accounts cover measurements at birth (length and weight), maximum measurements for males and females, and life span. The World Whale Watcher will appreciate the description of distinct flippers and fins. No identification key is given, but the many zoological details might even allow identifica- tion of dead whales from beach surveys. The well- designed and structured index allows for a quick identification in the field and on a moving boat. The species accounts are structured in four major groups of marine mammals: marine fissipeds, pinnipeds, cetaceans, and sirenians. Unfortunately, a table of contents naming each individual species is missing. But the illustrated glossary is very good since it covers so many details. The list of “regional sea mammal assemblages’”’ is rarely found elsewhere in a field guide. The World Whale Watcher gets really spoiled with this book due to the documentation of all sea mammals living in the Tropics, Rivers, Pelagic Oceans, Arctic, and Antarctic. The chapters about Fur Seals and Sea Lions are real beauties; the book allows the reader to compare freshwater seals like the endemic Caspian Seal and Baikal.Seal (hard to confuse them in the wild though). The chapter about the 19 Beaked Whales is another gem, even showing sharp photos for many of these rare creatures. Presenting 120 species accounts in a row might appear to be a little boring for the average reader. However, this book contributes greatly to global sea mammal conservation: one can only protect what one knows. Specifically the seal chapter emphasizes the importance of international legal frameworks like the Conservation of Antarctic Seals. The authors mention that the extinction of Gray Whales in the Atlantic is still a mystery. But on the positive side, a new species of sea mammal — the Pygmy Beaked Whale — was described as late as 1991. For com- pleteness, the extinct Steller’s Sea Cow, the Caribbean Monk Seal, and the Japanese Sea Lion are mentioned as well. The book emphasizes once more that the major threats to sea mammals stem from gillnet fishery, by-catch, aquaculture, hunting, global change, oil and gas development, and pollution. Some of these general perceived threats might suffer from “spatial biases” (locations where intense research studies were carried out). As a realistic “World Whale Watch” test, I put this book for 15 minutes under water: it hardly survived the true “at-sea” conditions; but at least it floats and it resists water spray! Perhaps, new whale watchers might find it confusing that Killer Whales and Pilot Whales are treated as “Ocean Dolphins” and that these species cannot be found in the “regular whale” sections. The complex Ocean Dolphin chapter I find somewhat “crowded”. It is too bad that none of the heavily debated population numbers mentioned in the species accounts carry any scientific references, which is crucial for the transparency of such state- ments in the public. It would likely be of great ser- vice to the informed user to make the distribution maps of this book digitally available. But really, that’s all “peanuts” and does not subtract from the value of this splendid publication. After reading this guide, one simply wants to buy a boat and see all these unique and fascinating creatures in the wild. FALK HUETTMANN Geography Department — Earth Science, 2500 University Drive N.W., University of Calgary, Calgary, Alberta T2N IN4 Canada, e-mail: falk @ucalgary.ca 2002 BOOK REVIEWS 505 Cetacean Societies: Field Studies of Dolphins and Whales Edited by J. Mann,, R. C. Connor, P. L. Tyack, and H. Whitehead. 2000. University of Chicago Press, Chicago. xiv + 435 pp., illus U.S. $35. “No longer must we kill whales to study them”. Instead, and as brilliantly presented in this book, innovative observational studies are nowadays car- ried out to learn about whales and dolphins (cetaceans). This book is indeed a cetacean landmark publication; it sets examples for many other species groups and will be of great value to the cetacean scholar as well as to the general reader interested in wildlife behaviour. However, it also applies approaches seen already in the research of apes. So is the “bible”, Primate Societies by Smuts et al. (1987), used as a role model for this book. This approach pre-defines automatically the philosophy and conservation focus. “Until recently, much of what we knew about cetaceans came from commercial sources such as whalers and trainers for dolphin shows”. The rela- tionship between scientists and members of the marine mammal public display community can be an uneasy one. Here we see a book based on many years of innovative field studies of wild populations. It is devoted to Ken Norris, a forefather of cetacean research. Thirteen world-wide renowned authors con- tributed to this volume, most of them are US-based and US-funded. The presented cetacean studies come from major cetacean study locations in the world, such as Sarasota (Florida/U.S.), Shark Bay (Western Australia), Galapagos Islands, Moray Firth (Scotland), Southwest Canada, Eastern Canada, and others. The book is conveniently divided into three parts: (i) History and Methods, (ii) Four Species (Bottlenose Dolphin, Killer Whale, Sperm Whale and Humpback Whale), (iii) Comparative Studies, Theory, and Conservation. The Appendix is very interesting as well and deals with phylogeny and tax- onomy of cetaceans. Very helpful for the reader is also the subject index and the citation index, which stimulates extensive use of the strong 57 page (!) ref- erence section. The book makes a stark claim for a science that is question-focused and not only taxon focused. “Whales and dolphins are long-lived, reproduce slowly, and spend most of their lives below the water’s surface”. Therefore, they are very hard to study. This book helps to overcome this problem by devoting a great chapter to all major (boat-based) sampling approaches used in behavioural studies of cetacean societies such as Focal Sampling, Scan Sampling, Rare Events, etc. In addition, relevant field sampling issues are outlined in detail in order to assure valid inferences from field observations. The scholar of cetaceans will appreciate as well the other specific and splendid chapters allowing for great information, such as the History of Studying Cetacean Societies, Identification by Photo, Tagging, Accoustic and Genetics. Of interest is also the topic about John Lilly and the “myths” that he created in the public mind. Once more, the Killer Whales of British Columbia, residents and transients, get cov- ered in full length. Right Whale studies are not directly included, but often referred to. “Research on cetacean societies is in its infancy compared with that on primates, carnivores, or ungulates”. All relevant subjects currently discussed in the literature for wildlife societies are presented in this book. It deals with such ‘hot topics’ like fission- fusion, migration, tool use, vocal learning, benefits of group living (mutualism, altruism, cooperation), roving males, mating strategies, Humpback lecks, parental investment, sperm competition, infanticide, philopatry, gender relationships (male-male, female- female and male-female), gambits of being in a group, communication (tactile, electric, chemical, communication, visual and acoustic), parasite load in Humpback whales, and effects of Global Change. Cetacean Societies is clearly an outstanding book in the field of behaviour studies, but, at the same time, it is lacking the presentation of real field datasets. The book primarily presents summarized results rather than any raw data (no relevant databas- es are reported, nor where and how they could be accessed for the public, e.g. via www). However, important (female) cetacean data on life-history, demography and many other subjects are compiled in over 20 tables; e.g., studies on travel speed. In addition, many fascinating photos taken from field work are presented in the 16 colour plates and 93 half-tones; 41 line drawings are included to show graphs and to visualize concepts. As it may not be missing in any popular book dealing with cetaceans (but not the prime focus of this book) interesting chapters about Present Exploitation, Conservation and Whaling are includ- ed. These chapters point out that Aboriginals over- hunt cetaceans in some locations in Canada. Whaling was developed from a fisheries mentality; so it was based on pure exploitation without considerations of a sound sustainability. No wonder that according to some sources up to 95% of the Humpback pre- exploitation population was eliminated. Japan’s Striped Dolphin culls appear not to be sustainable either. Rather than cetaceans competing with fish- eries, there can be absolutely no doubt that “cetaceans are more threatened by fisheries than the reverse’. The IWC (International Whaling Commission), a major player when it comes to cetacean conservation and research, gets only mentioned in brief detail. But the book cites that “whale-watching around the world was estimated to be about 500 Million US$ in Tt _—— MEN I Se WE Sige us 506 1994, roughly equivalent to 25,000 minke whale car- casses”. The authors report that the effects of Whale Watching tourism on Killer Whales and others are unclear and still deserve more research. The reader will find the overview table about cetaceans’ conservation status and IUCN categories of great value. By-catch, habitat degradation and hunting are among the 16 prime threats for the con- servation of cetaceans. The hot subject of “by-catch’ is not covered in much detail, but at least it is men- tioned. Dams in rivers can interrupt and fragment sea mammal populations. Military exercises and indus- trial blastings; e.g., from oil explorations, are well known to be harmful as well. “Until quite recently, oceans and even rivers, were perceived as almost infinite sinks that could absorb human waste with no impact on humans and little harm to the environment.” How true. The MARPOL convention is supposed to fight marine pollution caused for instance by plastic, chemicals, oil or sewage (still, none of the coastal Canadian cities have a sewage system worth mentioning). The book emphasizes nicely that habitat degradation is the major component threatening the entire marine ecosystem (two thirds of the world) and that there are no administrative and national boundaries to pol- lution, nor to whales (animals move beyond borders and don’t care about national and administrative units). The authors present a chapter on Conservation, Protection and Science which shows how scientists can contribute to this important task. Besides giving a perfect overview and summary about current cetacean research topics, this book also BOTANY Lichens of North America By Irwin M. Brodo, Sylvia Duran Sharnoff, and Stephen Sharnoff. 2001. Yale University Press, New Haven. 795 pp., illus. U.S.$69.95 Exquisite, aesthetically pleasing, wonderful, breath-taking, and magnificent barely describe the 32 full-page color pictures and nearly 900 hundred smaller ones in this book. Although they alone are worth the price, there is much more in this book. A total of 804 species are illustrated plus 250 other common North American lichens are included in the keys. These species represent about 30% of the lichens in North America. The book has two major parts. “About the Lichens” occupies the initial 113 pages, and is com- posed of 14 chapters that enumerate lichen features, biology, including discussion of the symbiotic rela- tionship between fungi and algae that make the THE CANADIAN FIELD-NATURALIST Vol. 116 shows what still needs to be researched. For instance, the book points out that nighttime research on killer whales does not exist. Behaviour approach- es to systematics and taxonomy could be useful as well; so could be many aspects of behaviour-based predator-prey studies. For my personal taste, too many of the questions asked and addressed in this book deal with “invest- ment” and energy effort, somehow resembling pure money and banking issues, and making cost and ego- istic gain considerations rule. Our behaviour research is ill-fated if only the bank/money approach, which humans face in their daily life nowadays, gets com- pletely transposed on animals and behaviour interpre- tation; is that all what the science of animal behaviour currently can offer? It simply presents a limited view, and a plain copy of approaches applied to monkeys, and used earlier by Nobel-prize winning Gary Becker ‘The economic approach to human behaviour’. Unfortunately, results and contributions from “sci- entific whaling” are not included in this book. Might be there are none; time will tell. Proceeds from this affordable volume go towards a fund to support graduate students on behaviour studies. This mile- stone reference is a grace to review and to read. Every naturalist and wildlife scientist should have it in her/his bookshelf. FALK HUETTMANN Killam Fellow, Geography Department—Earth Science, 2500 University Drive N.W., University of Calgary, Calgary, Alberta T2N 1N4 Canada; e-mail falk @ucalgary.ca lichens, colors, chemistry, habitats, distribution, and classification. Additional chapters treat lichens and people (food, clothing, dyes, perfumes, medicines and poisons, human impact on lichens, and much more), environmental monitoring with lichens, col- lecting and studying lichens, and how to use this book to name a lichen. The text is supplemented with photographs and drawings to illustrate salient points. The writing is careful and deliberate, so it does not scare off newcomers to the lichens. Con- cepts, features, and methods are explained simply but in detail. As with any specialized area there is terminology to be grasped. Words that may not be familiar to naturalists are explained in the text where they first appear and the glossary is also an aid to understanding terms by giving a description and often a reference to a figure. 2002 “Guide to the Lichens” begins with a 27-page key to the identification of lichen genera and major groups. The use of a “key” is explained in chapter 14. The largest part of the book is a chapter, of 604 pages, titled “Description, illustrations, keys to species, and maps”. The lichens are arranged alpha- betically by genus. Thus if you know the name of the lichen it is easy to find its description. I like the alphabetical arrangement but others would prefer a grouping of similar types (i.e., all the orange species together or all the foliose species grouped), such groupings are used in the keys. The treatment of each lichen is composed of sev- eral parts. In addition to the color photograph, there is the scientific name and, for many, a common name. The essential features, concentrating on the macro characters, of the lichen are enumerated as is the habitat, but habitat comments are brief, probably due to space limitations. For example, Solorina cro- cea in south central British Columbia is common in dry lodgepole pine forests on sandy soil, whereas the text states “usually in moist spots ... or seepage areas in arctic or alpine sites.” The geographic distribution of each species is shown on a map. Maps fascinate me. And they convey a consider- able amount of information. The maps quickly tell the user whether the lichen is found in one’s neigh- borhood. Disjunct distributions can pose intriguing questions. Why does Parmotrema stuppeum occur only in two areas that have little apparent similarity, the Appalachian Mountains and a narrow band along coastal California? Current distributions can reflect earlier phenomena. Umbilicaria caroliniana’s occur- rence in only two Ice Age refugia, in the southern Appalachians, and northern Alaska and the Yukon, suggests it was widespread in pre-Ice Age times. Other species, like Peltigera aphthosa, are wide- spread and their distribution may reflect its ability to compete effectively with other plants. Reviewers routinely trumpet the glories of the subject but they should also aid the users by indicat- ing difficulties. There is little to complain about in Lichens of North America, and then the items are quite minor and do not distract from the quality and usefulness of the volume. When I first opened the book, I was struck by the beauty of some full-page and smaller photographs (for example, pages ii—xvi, 114, 116) and wondered which lichen I was admir- ing for these pictures were not labeled. The Cup BOOK REVIEWS 507 Fungus (page 4) although labeled Aleuria aurantia has the morphology and habitat of Sowerbella rhenana. The distribution maps show all of Canada and the United States, except Hawaii. Most are use- ful, but lichens of restricted distribution are uncer- tainly discernable on the map (e.g., Lecanora phry- ganitis and Niebla combeoides). For these species arrows indicating their location or a restricted map (i.e., only of the southwest) would have been more informative. Figure 533 is slightly out of focus in my copy of the book. Brodo has had a long association with The Ottawa Field-Naturalists’ Club and its journal The Canadian Field-Naturalist. The OFNC can be proud of its sup- port of the publication of Lichens of North America. And the other benefactors are to be thanked for fore- sight in supporting this project. One direct result of their participation is the high quality of the printing and binding and the book’s relatively low price! This book is the end product of an enormous pro- ject that took years of devotion to complete. Brodo writes (page xx) “the road to a finished manuscript was a rocky one,...” And one rock, reported by col- leagues in Ottawa, was when managers at Canadian Museum of Nature, where Brodo spent most of his career, at one period near its completion prohibited work on the project. Brodo persevered at home dur- ing evenings and weekends; however, for support at the end, the title page graciously states, “published in collaboration with the Canadian Museum of Nature,” and the CMN is also fairly acknowledged for its career support of Brodo’s fieldwork and travel. In summary, the book is highly recommended. The style of writing, the superb photographs, and the reasonable cost are unbeatable. A glimpse at this book should be sufficient to convince naturalists, ecologists, and others to get involved with this fasci- nating group of organisms. Some of the photographs in this book can be seen at www.lichen.com where one can also find more background on lichens, the preparation of the book, and the authors and their interests. Regrettably, Sylvia Sharnoff did not live to see the final publication. J. GINNS 1970 Sutherland Road, Penticton, British Columbia V2A 8T8 Canada 508 MISCELLANEOUS THE CANADIAN FIELD-NATURALIST | : Vol. 116 Life in the Treetops: Adventures of a Woman in Field Biology By Margaret D. Lowman. 1999. Yale University Press, New Haven. xvi + 219 pp., illus. U.S. $27.50. During the past two decades an increasing number of books, essay collections, biographies, and autobi- ographies have dealt with the lives, successes, and tribulation of women scientists. Most have focussed on women in the laboratory sciences, and only a few detailed the challenges women faced when undertak- ing fieldwork. Margaret Lowman’s autobiography takes us in a different direction — upwards to the canopies of Australian, American, and African forests. In addition to its scientific aspects, this auto- biography also presents a vivid picture of gender relations in the Australian outback. As readers of the Canadian Field-Naturalist know, field research does not follow office hours. Biologists in the field do not work seven hours per day, five days a week with well defined rest periods and vacation times. Birds, plants, butterflies and other organisms have life cycles that do not conform to human biology and late twentieth century lifestyles. The exigencies of fieldwork do not match the rhythms of marriage and parenthood. Life in the Treetops provides a lively personal account of the successes and setbacks of an American woman field biologist on three different continents. Born and educated in New England, Meg Lowman enrolled in graduate school in Scotland, completed her Master’s degree on “the seasonality of highland birch trees” (page 10) and, influenced by her advisor’s interest and enthusiasm, decided to do her doctoral research on the tropical rain forest. She chose Australia because it was both “English speak- ing...[and its] forests remained some of the least studied in the world” (page 11). With a scholarship from the University of Sydney botany department she soon enjoyed the scientific opportunities provid- ed by this continent’s forest ecosystems, but was unaware of the antiquated attitudes to women and conservation awaiting her. The author’s account of her scientific fieldwork in the treetops is both fascinating and amusing. She provides sufficient information for the uninitiated to understand the scientific questions she had asked and her varied methods of canopy research. She does not minimize the dangers, inconveniences, and difficul- ties of this sort of research for any one, particularly for a woman. Climbing to the canopy level using a rope, or being lifted to the treetops by a cherry pick- er, a construction crane, or a hot air balloon equipped with a canopy sled (towed under the bal- loon), is not for the fainthearted. Neither is building and using a canopy walkway. And these were only the technical challenges than enabled Meg Lowman, her colleagues, students, and friends to get to their field sites. Inclement weather, insect pests, poi- sonous snakes, and frightened Brush Turkeys were all parts of the environments she studied: Her vari- ous projects included the “growth patterns on rain forest leaves,” studies “of insect herbivores in action,” “plant-insect relationship,” “dieback” of trees which led to challenges in applied ecology, as well as “seedling lottery” at ground level (page 94). Accounts of the author’s personal life are well integrated into the book. We learn about her attempts to juggle the roles of scientist, wife, mother, and homemaker in a social environment that did not vali- date or even understand her scientific interests and aspirations. This part of her story is not unusual, but is rarely documented. Women scientists of earlier periods, with the exception of Margaret Morse Nice, did not disclose personal relationships and private stumbling blocks that often delayed, slowed down, or even prohibited their scientific activities. We rarely read about taking infants in diapers into the field, disapproving mothers-in-law, or a restrictive environment that would prompt a scientist to read the Journal of Ecology “wedged between the pages of ... Women’s Weekly” (page 103). Throughout her studies and professional life Meg Lowman was encouraged and supported by her par- ents, teachers, advisors, and colleagues. She missed having a female mentor in Australia and did not seem to meet women botanists. While the University of Sydney may not have employed women in the biolo- gy department at the time, there were many women botanists and other scientists in Australia. Among them was Gretna Weste of the University of Melbourne who also studied dieback of the native forest and grappled with issues of motherhood and scientific research a generation earlier. Clearly, Meg Lowman was not yet part of the “intellectual spider- web...that interconnects” scientists (page 136). In time, as an Australian researcher, teacher at Williams College in the U.S. and, since 1992, Director of research and conservation at the Marie Selby Botanical Gardens in Sarasota, Florida, she devel- oped her own worldwide network of scientists. This well written and delightfully illustrated book ought be read by seasoned and aspiring scientists alike, as well as the general public. It should be on the bookshelves of public libraries, high schools, community colleges, and universities. I hope that instructors in biology, history, and women’s studies will take note of this slim but important volume and use it in their classes. I know that I will. MARIANNE GOSZTONYI AINLEY Professor, Women’s Studies/History, University of Northern British Columbia, Prince George, British Columbia V2N 4Z9 Canada 2002 BOOK REVIEWS 509 Victorian Sensations: The Extraordinary Publication, Reception, and Secret Authorship of Vestiges of the Natural History of Creation By James A. Secord. 2000. The University of Chicago Press, Chicago. 624 pp., illus. U.S.$35. Should we be curious, after 150 years, about a little book titled Vestiges of the Natural History of Creation? Originally published in 1844, 15 years before Charles Darwin’s On the Origin of Species, it prefigured the latter by popularizing the idea of evolu- tion. Yet, why is it of any importance? Is it the fact that it was one of the early works to attempt to reach the general reading public rather than the more scien- tific elite; the fact that it was anonymously written; or is it the time and place it was published in conjunction with the political, social, or religious views of the day? James A. Secord, editor of the recently reprinted version of Vestiges (University of Chicago Press, Chicago, 1994) asks these questions. I found this work wonderful if only for the fact that unlike the few other history-of-the-book projects, Victorian Sensations puts Vestiges in a multitude of contexts. Seemingly unrelated, they all interconnected in skeletal form with each other. And not just the obvious social, political, or religious arenas, but even with the art of making the book, the marketability from an author and publisher’s point of view. Reaching a broader audience without appearing as a ragish tome was a major concern. The author of the scandalous work, a work that combined all manners of science, was publisher Robert Chambers (1802-1871). But to avoid any link to his good, pros- perous name, his manuscript was issued with another publisher. Through a third party, both author and pub- lisher masterminded the physical publication (and the numerous editions to follow), and even selected some of the reviewers. The work contained snippets from many scientific disciplines and was formulated for the general reader in a kind yet somehow authoritative manner. From the formation of the solar system, the fossil record, to humanity, all under the guiding principle of some sort of evolutionary ideology. Conveying such a broad sci- entific spectrum, especially when many of these sci- ences were still in their infancy, was unattainable and thus not without errors. This provided much fuel to the cynical, put-off reviewers and many of the general public. As one scientist wrote, “It is humiliating to think that a work of such flimsy pretensions should have obtained so much consideration, because it proves how little the intelligent public are aware of the real progress of science” (page 212). But it placed evolutionary thinking in the public consciousness like it never before had. Even if we can’t fathom the reasoning behind the historic place of Vestiges, we surely must understand, or at least appreciated what it did for Charles Darwin (1809-1882). For Darwin, it was not the contents of the book that mattered, but the manner in which the public and professional reader received it. As Secord notes, “Reading reactions to Vestiges became a way [for Darwin] of imagining the reception of his own, unpublished essay” (page 431). Secord goes on to substantiate his claim that though culture and society has raised Darwin’s stature above all but a few in the scientific arena, as no one would disclaim the impact of The Origin of Species, the danger of supporting the mythology the “Darwinian” Revolution can be rather myopic. To the general reader “It obliterates decades of [unheralded] labor by theologians, technicians, printers, editors, and other researchers, whose work has made evolutionary debate so significant during the past two centuries” (page 518). Works like Vestiges. The reflective nature of Victorian Sensations, the broad spectrum of its contexts, and the resources upon which it is supported are commendable. And for those who dismiss the idea of a work’s place in time, its construction, and mode of dissemination, this title is definitely worth a look. TIM T. TOKARYK Box 163, Eastend, Saskatchewan SON OTO Canada Northern Wild, Best Contemporary Canadian Nature Writing Edited by David R. Boyd. 2001. Greystone Books, Douglas & McIntyre, Vancouver. 278 pp. $22.95. Someone has finally done it. David R. Boyd, a Senior Associate with the Eco-Research Chair in Environmental Law and Policy, University of Vic- toria, has put together one of the surprisingly few anthologies of nature writing by Canadians — in con- trast to the scores of U.S. nature writing anthologies published over the years. Northern Wild, an attempt to fill that void, includes a wide variety of contemporary texts on topics as diverse as slugs, berry patches, Inuit navigation, home, bugs, the prairie landscape, fishing, and more (in fact, I’d wager that the editor is a fisherman, since three of the 20 texts are definitely about fishing). They range from shorter to longer, lighter to heavier, more person- al to more scientific, more humorous to more serious pieces — by authors both lesser- and better-known. Different as the essays are, they all share, as Boyd writes in his introduction, “something that we all enjoy as children but tend to lose as we grow up” — a sense of wonder. 510 It would be best to let some of the writers speak for themselves. John B. Theberge experiences wonder and truth in the gaze of a wolf waking from tranquilization: “She held her gaze for a full minute, two minutes, an eter- nity. Wolf — human — each searching for meaning in the eyes of the other. Eyes so alike — iris, pupil, cornea, lens, size, musculature, movement. But eyes so different, reflecting two different social orders that began diverging, like two continents drifting apart, hundreds of thousands of years ago.” Beth Powning finds wonder and peace in her own home: “The comfortable, homey chirp of swallows and the snipes’ mournful, swooping wing-whistle are the sounds of early morning, familiar as water filling a kettle ... | smell the dry-wood, old farmhouse smell of wood smoke and mice; and daisies, their bitter scent riding a dancing breeze ... I can’t imagine ever leaving this place.” Des Kennedy shares the wonder and fascination of a slug crawling on his hand: “After a few moments of frightened withdrawal, its head emerged, the THE CANADIAN FIELD-NATURALIST TE i mini iiha © | Vol. 116 - slender optic tentacles extending like time-lapse flowers. As it began to sense the world around it, one — then both — of its smaller tentacles touched the surface of my skin delicately. I watched the animal apply its pedal gland to my palm. Attaching a silvery ribbon of mucus to my skin, it slipped forward — a cold, smooth softness on my hand.” Heather Menzies feels wonder and meaning in the company of a pine she helped plant as a child: “Around me, shadows drift up like root hairs seeking passage. A breeze whiffles the upper branches, light as surf bubbles caressing a shore. I sit on the moss, enveloped by these trees I helped grow into this ground over so many years. They’re in my blood now. I feel them.” That’s only a brief sampling of the fine writing and diversity of topics represented in Northern Wild. It’s a delightful and meaningful read. R. SANDER-REGIER RR5 Shawville, Quebec JOX 2Y0O Canada Cheltenham in Antarctica: The Life of Edward Wilson By D. Wilson and D. Elder. 2002. Reardon Publishing, Cheltenham, England. 143 pp., illus. £9.99. Edward Adrian Wilson was born into Victorian comfort and anonymity. He died in appalling need and with international acclaim. This remarkable man achieved more in his short life than most of us could in three lifetimes. As a teenager I read the accounts of Scott’s Antarctic expeditions, which included Wilson, and compared them to Shackleton. I decided that Scott fared poorly when compared to Shackleton. Scott was part of the gentry and followed the British class system. Shackleton was a team player. Scott died, as did many of his crew. Shackleton lived and no man under his direct command ever died. I always felt it was unfair that Scott’s men were more famous heroes in their day than Shackleton’s group. To me Scott’s efforts missed the mark because he did not take common-sense advice, whereas Shackleton was a success. I have softened this stance over the years, as I understood more about the contribution made to science by Scott’s team. Despite my unfriendly view of Scott I have always been a fan of Wilson. After all he was the naturalist and artist. True, his books have a “jolly hockey sticks” flavour that comes from his privi- leged upbringing. Yet he did so many interesting things. This new biography chronicles the entire life and times of this extraordinary man. The authors, Wilson (a relative) and Elder, have put together a smooth flowing narrative that captures the nature of Ted Wilson’s spirit as well as his history. Some biogra- phers include so many quotes of their subject’s own writings that the rhythm of the text is destroyed. These authors have used Wilson’s quotes but have done so with discretion. The quotes add extra under- standing of Wilson’s character without being disrup- tive. For a book of its size — it is as big as a Reader’s Digest magazine — it has a large number of illustra- tions. Many of these are black-and-white family and expedition photographs. In addition there are a num- ber of colour reproductions of Wilson’s artwork. Remarkably some of these date back to his child- hood, so you can see his progress to a full-fledged Victorian nature artist. A number of the watercolours are in colour. As well as being delightful paintings many have a historical significance too. I have long been puzzled by Wilson’s death. The classic version is the he, Scott and Bowers were trapped in their tent by a 10-day blizzard and died of starvation. But Wilson, with Cherry-Garrard and Bowers, made a 5-week trek to an Emperor Penguin nesting colony in the middle of winter. This journey was made under appalling conditions (read The Worst Journey in the World by Apsley Cherry- Garrard) and they all survived. How then could these seasoned travelers, who had braved blizzards before, wait for death when salvation was only 11 miles away? Recently I read that Scott had allowed 4300 calories per day, with more protein than fat. Recent studies have shown a person can in a severely cold 2002 climate burn 11 000 calories per day, and needs more fat than protein. These men had been starving for weeks, and starvation affects the brain’s logic as well as the body. Maybe they were disoriented and confused. A second intriguing theory notes that there is no other record of a 10-day blizzard in this region. The longest was four days and this fits with the physics of the terrain. Did Wilson tell Scott, who was suffering badly — probably dying — from frost- bite, that there was a blizzard so he would not know they had chosen to stay with him even though it meant their own death? This would fit with Wilson’s character and deeply held principles. A most unusual addition to this biography is a guide to town and country walks in Wilson’s home neighbourhood. These direct the reader to the loca- tions Wilson knew and visited, and to locations of historical significance. This is a clever and charming bonus. BOOK REVIEWS 511 This is a delightful book, recounting the life of a most remarkable man in an extraordinary period of history. His work had a profound impact on science; something that is often forgotten in the face of his heroic death. But there is something else about him and so perhaps the best note to end with is that his contemporaries said, “everybody loved him.” After reading this book, I am sure I would have too. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6KS5 Addenda: Just as I was finishing this review I discovered that Random House has published Cherry: A Life of Apsley Cherry-Garrard, by Sara Wheeler (See paragraph six above). After years of anonymity there is suddenly a won- derful revival of interest in the golden age of Antarctic exploration. th ——— EEE i hia ~§ | - News and Comment Canadian Species at Risk May 2002 Issued by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC), the list is 34 pages contain- ing: About COSEWIC, (mandate, membership, definitions); — Summary Tables (COSEWIC species at risk, not at risk, and data deficient; results of May 2002 COSEWIC meeting), COSEWIC Lists (Explanation of symbols, Geographical occurrence and abbreviations; List 1 Species designated in the five “risk” categories, List 2. Species examined and des- ignated in the Not AT RIsK category; List 3 Species exam- ined and designated in the DATA DEFICIENT category) — Record of Status Re-examinations — List of name changes — COSEWIC Species Specialist and Aboriginal Traditional Knowledge Subcommittee Contacts (2002-2003). It is available from COSEWIC Secretariat, c/o Canadian Wildlife Service, Environment Canada, Ottawa, Ontario K1A 0H3. See Web site: http://www.cosewic.gc.ca Froglog: Newsletter of the Declining Amphibian Populations Task Force (51, 52) Number 51, June 2002. Contents: The DAPTF in Africa (Tim Halliday) Chytridiomycosis Survey in South Africa (Che Weidon); Amphibian Faunal Diversity and Conservation Status in The Niger Delta Basin (Southern Nigeria): A Update (Godfrey C. Akani and Luca Luiselli); Amphibian Research in Myanmar (Guin Wogan and George Zug); South Asian Amphibian CAMP; Froglog Shorts; Publications of Interest. Number 52, August 2002. Contents: Northern red- legged frpgs and emdocrine disrupting compounds (EDCs) (James B. Bettaso, Hartwell H. Welsh, Jr., and Brent D. Palmer; Monitoring amphibians and reptiles along the Drava River (Tibor Kovacs); South Asian Amphibian Marine Turtle Newsletter (97) July 2002. Contents: Articles: Cold Stunned Loggerhead Turtles in South Adriatic Sea; Use of Pop-Up Satellite Archival Tags to Quantify Mortality of Marine Turtles Incidentally Captured in Longline Fishing Gear; Satellite Tracking of Post-Nesting Movements of Green Turtles Chelonia mydas from the Gangkou Sea Turtle National Nature Reserve, China, 2001; Notes: Happenstance or Design: An Unusual Association between a Sea Turtle, Octocoral and Barnacle; Apparent Beach Basking of an Atlantic Green Turtle (Chelonia mydas) at Dry Tortugas National Park, Florida; A Record of the Northernmost Verified Leatherback Sea Turtle Nesting Event on the East Coast of the USA; Leatherback Turtles in the Mid-South Atlantic Waters; MEETING REPORTS; BOOK REVIEW; Workshops: Executive Summary; Froglog Shorts; Publications of Interest. Froglog is the bi-monthly newslett;er of the Declining Amphibian Populations Task Force of The World Conservation Union (IUCN)/Species Survival Commission (SSC) and is supported by The Open University, The World Congress of Herpetology, and Harvard University. The newsletter is Edited by John W. Wilkinson, Department of Biological Sciences, The Open University, Walton Hall, Milton Keynes, MK7 6AA, United Kingdom; e-mail: daptf@open.ac.uk. Funding for Froglog is under- written by the Detroit Zoological Institute, P. O. Box 39, Michigan 48068-0039, USA. ANNOUNCEMENTS; NEwS & LEGAL BRIEFS; RECENT PUBLICATIONS. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Research Group, School of Biological Sciences, University of Wales, Swansea, SA2 8PP Wales, United Kingdom; e-mail MTN @swan.ac.uk; Fax +44 1792 295447. Subscriptions to the MTN and donations towards the production of MTN and its Spanish edition NTM [Noticiero de Tortugas Marinas] should be sent to Marine Turtle Newsletter c/o Chelonian Research Foundation, 168 Goodrich Street, Lunenburg, Massachusetts 01462 USA; e-mail RhodinCRF @aol.com; fax + 1 978 582 6279. MTN web- site is: . S12 2002 Point Pelee Natural History News 2(2) The summer 2002 issue contains: ARTICLES: The Tiger Beetles (Cincindelidae) of Point Pelee National Park, Ontario (Stephen A. Marshall) — Noteworthy Bird Records: March to May 2002 (Alan Wormington) — Chuck-will’s-widow: Second Record for Point Pelee, Third for Ontario (Robert Curry) — In Search of a Chuck-will’s- widow Nest (George D. Bryant) — Chuck-will’s-widow: A Nest is Found! (Alan Wormington) — The Decline of Grassland Birds: Some Insights and Comments (Alan Wormington) — IN THE FIELD — NEWS AND ANNOUNCE- MENTS — UPCOMING EVENTS AND OUTINGS. THE OTTAWA FIELD-NATURALISTS’ CLUB AWARDS 513 This newsletter for Point Pelee National Park, Ontario, is edited by Alan Wormington (e-mail: wormington» @juno.com). Editorial Assistants are M. Lea Martell and Matthew J. Smith. The web site is www.wincom.net /~fopp/Natural_History_News.htm. Subscription rates are Canada: CAN $15 (one year) or $30 (two years); International: US $15 (one year) or $30 (two years). Send payment (and e-mail address, optional) to The Friends of Point Pelee, 1118 Point Pelee Drive, Leamington, Ontario N8H 3V4 Canada. Issues are mailed in March, June, September, and December, and back issues are available for $15 per Volume/ $5 per issue (postage paid). Recovery: An Endangered Species Newsletter (21) June 2002 This issue contains: recovery highlights: New project, Restoring habitat, Conserving frogs; profile: Biologist champions ecosystem recovery [Marilyn Fuchs] (David Wylynko); News BITES: Custom-designing recovery; Conserving shrike habitat; Protecting a Quebec plant; Butterflies at risk; COSEWIC Update; CITES Update; RENEW Update; ESRF update; FIELD NOTES: Program asisits reptiles at risk; Monitoring whales; Falcon a celebri- ty; ANNOUNCEMENTS: Awards; New publications; Upcoming event; Site seeing; FEATURED SPECIES: Recovering the Barn Owl (Bernt Solymar). Contact: Recovery, Canadian Wildlife Service, Environment Canada, Ottawa, Ontario K1A 0H3. Web site: www.cws- scf.ec.gc/recovery/archive.html Minutes of the 123" Annual Business Meeting of The Ottawa Field-Naturalists’ Club, 8 January 2002 Place and time: Chairperson: Attendance: Eleanor Zurbrigg, President. Thirty persons attended the meeting. Victoria Memorial Building, Canadian Museum of Nature, Ottawa, Ontario, at 7:30 pm. Attendees spent the first half-hour reviewing the minutes of the previous meeting, the Treasurer’s report and the Report of Council. The meeting was called to order at 8:00 pm. 1. Minutes of the Previous Meeting It was moved by Dave Moore and seconded by Fenja Brodo, that the minutes be accepted as amended. (Motion Carried) 2. Business Arising from the Minutes There was no business arising from the minutes. 3. Communications Relating to the Annual Business Meeting There were no communications relating to the Annual Business Meeting. 4. Treasurer’s Report Frank Pope reviewed the financial report for the year ending September 30, 2001, noting that the Club’s net assets had increased by $37,584. Most of this increase was the result of a surplus of $25,895 in the fund allocated for the CEFN. This is due to falling behind in the publishing schedule as well as not hav- ing to purchase paper for the CEN this fiscal year. Moved by Frank Pope and seconded by Roy John that the Financial Report be accepted. (Motion Carried) 5. Committee Reports Eleanor Zurbrigg introduced each of the Committee reports and a representative of the appro- priate Committee and asked for questions and comments. Amendments were made to the reports of the Birds, Computer and Finance Committees. Dave Moore asked if we had received any feed- back about the materials that had been supplied for the presentation by Tom Bearss to the Trinidad Naturalists’ group. Eleanor Zurbrigg said that we had heard that it had gone well. Frank Pope asked if we still received their annual report. No one was certain of this, but we do still get their newsletters. Moved by Dave Hobden, seconded by Bev McBride, that the reports as amended be accepted. (Motion Carried) 6. Nomination of the Auditor Moved by Frank Pope, seconded by Roy John, that Janet Gehr continue as Auditor for another year. (Motion Carried) 7. Report of the Nominating Committee The Committee recommended the following list of candidates for the 2001 Council (new members are indicated by an asterisk) President: Eleanor Zurbrigg Vice-President: Roy John Vice-President: Gary McNulty Recording Secretary: Ken Allison Treasurer: Frank Pope Other Members Irwin Brodo Rod Bedford Bev McBride Bill Cody Bob Roach Francis Cook Stan Rosenbaum Karen McLachlan- John Cameron Hamilton Roy John Dave Moore Peter Goddard* Charlie Clifford* Louise Schwartz* Kathy Conlan* David Hobden Diane Holmes* Barbara Gaertner Diane Lepage* Dave Smythe Four members of the 2001 Council decided not to stand for re-election: Tony Halliday, Terry Higgins, Rita Morbia, and Fenja Brodo. Colin Gaskell thanked these members for their contribution to the Club. Moved by Colin Gaskell, seconded by Irwin Brodo, that the list of nominations for the 2001 Council be accepted. (Motion Carried) 8. New Business Fenja Brodo would like to see more articles from the committees in T&L. She feels that this would generate more interest in committees. Ernie Brodo is looking for new ideas for the Soirée. He feels that interest and attendance are declining. Pearl Peterkin responded that a group of public relations students from Carleton University have produced a communication plan for the club. One of their suggestions was to remodel the Soirée. Ernie also stated that natural history education needs increased visibility in our community. As a 514 2002 club we need to help generate more interest among young people. One suggestion was to provide an education award for teachers who showed out- standing achievement in the area of natural history education. Pearl Peterkin suggested it is time to change the award for the Science Fair. There was considerable discussion about this topic. Pearl ended the discus- sion by stating that any changes would now have to wait for 2003. Ernie Brodo suggested that we should attend teachers’ professional development days to offer our resources to strengthen natural history education. Frank Pope noted that this is being done, but that there has been very little concrete response in the end. Ruth Allison pointed out that any material would have to be targeted to the curriculum. Frank said that the FON provides this sort of targeted MINUTES OF 123RD ANNUAL BUSINESS MEETING 515 resources. Gary McNulty suggested that club mem- bers get involved in presenting natural history sub- jects to young people. Dave Hobden asked if we could use the club website to showcase this sort of information. 9. Presentation by Francis Cook Francis Cook gave an interesting retrospective view of his experiences over many years as editor of The Canadian Field-Naturalist. 10. Adjournment Moved by Fenja Brodo, seconded by Gary McNulty that the meeting be adjourned at 9:40 pm. (Motion Carried) KEN ALLISON Recording Secretary The Ottawa Field-Naturalists’ Club Comittee Reports for 2001 Awards Committee The following awards for the year 2000 were presented at the OFNC’s Annual Soirée, which took place April 27, 2001. HONORARY MEMBERS Frank Pope: For his extraordinary and continuing service to the club in almost every capacity. Donald M. Britton: One of Canada’s most distinguished botanists, with very significant contributions to our knowl- edge of the Canadian fern flora. John (Jack) M. Gillett: For both his numerous contribu- tions to Canadian botany as well as his many years of ser- vice to the OFNC, most recently in his series of articles on the flora of the Ottawa region published in Trail & Landscape (T&L). Joyce and Allan Reddoch: For their years of service to the OFNC as council members, editor of T&L, articles in T&L, work on conservation matters, and for their contributions to our knowledge of the orchids of the Ottawa region. MEMBER OF THE YEAR Sandra Garland: For her work in producing a Club web- page outstanding for its design, ease of use and accuracy, and conducting maintenance on the site to keep it both fresh and timely. GEORGE MCGEE SERVICE AWARD Philip Martin: For his many years of service to the Excursions and Lectures Committee both as Chairperson and especially as field trip leader; also his activity in conservation matters, most recently regarding the Leitrim wetlands. ANNE HANES NATURAL History AWARD Robert E. Lee: Principally for his studies of frog popula- tions and his excellent article on the subject in Trail &Landscape, but also for his on-going observations on the natural history of the district. CONSERVATION AWARD - MEMBER Albert Dugal: For his on-going and persistent attempts to secure protection for endangered spaces in the Ottawa region, especially the Leitrim Wetlands and South Gloucester Conservation Area, over a number of years. CONSERVATION AWARD - NON-MEMBER David Miller, Planner, Region of Ottawa-Carleton: For his significant role in securing protection for a large area of the South March Highlands, his leadership in the Region’s Natural Environment Systems Strategy project (NESS), his input into the Regional Official Plan, identifying areas of environmental concern, and many other activities that have helped, and will continue to help preserve the natural envi- ronment in the Ottawa District. I. BRODO Birds Committee The Birds Committee participated jointly with the Club des Ornithologues de 1’Outaouais to run a successful Christmas Bird Count at the end of 2000. Plans are in place for the 2001 count. We also had a well-attended fall bird count in October 2001. We ran the Peregrine Falcon Watch at the downtown Ottawa nest site, which attracted many volunteers and club members. This year’s watch was short lived because the young birds died in collisions with a building. We anticipate running a watch in 2002. The Bird Records Sub-committee met several times during the year to review records of rare birds for the area. Results of their work will appear in an upcoming Trail & Landscape. The first year of surveying for the Ontario Breeding Bird Atlas 2nd edition was a great success with almost all our region’s squares being covered by an eager team of volunteers. Our seed-a-thon raised around $1000.00 for the Club’s bird feeders, which continue to be popular for both birds and 516 humans. Our bird study group held several sessions during the year which were well-attended. We continue to operate the rare bird alert telephone tree and the Ottawa Bird Status Line, which is a recorded telephone message providing cur- rent bird sightings. Two Birds Committee members were featured in a profile of the Ottawa Field-Naturalists’ Club in the newspaper of the Peace and Environment Resource Centre. B. McBRIDE Computer Management Committee The CMC met once in 2001 and identified and per- formed the following activities: 1. Activity: Since the CMC has been dormant for a few years previous files and information needed to be recovered and filed. Result: The files were recovered and with the help of Sandra Garland were filed at the Fletcher Wildlife Garden Office. 2. Activity: Recover and update the CMC inventory list. Result: Alan German recovered and confirmed the inven- tory list of 14 May 1999. 3. Activity: Upgrade or replace the Club Bookeeper’s Computer Background: Bookeeping is one of the clubs core functions and the existing computer was becoming obsolete and uneconomical to upgrade. The lack of e-mail capability was also a requirement that could not easily be met by the exist- ing computer. Result: The club recieved a donation of a P100 computer which with the expertise of Roger Camm was outfitted with all necessary equipment to allow it to function. The com- puter was installed at its final location and is now operational. Thanks to Roger Camm and Alan German. 4. Activity: Membership List Software Upgrade. Background: The current membership list is in dbase for- mat, which is becoming difficult to maintain. Result: The database can now be used and modified using a contact management product called Goldmine. Contact management software provides not only easier user inter- face, but also a path towards such capability as email mem- bership renewals. It was noted that the club membership mailing costs are about $500 annually. A test was designed to show that data records can be exported to Excel and imported into Goldmine. The conversion of all records to Goldmine is ongoing and has taken longer than forecast. B. ROACH Conservation Committee The main activities were in connection with the following: Richmond Conservation Area (RCA): Robert Marler was nominated to represent OFNC on the advisory committee (RCAMC) which now reports to the City of Ottawa. All nominations and the draft Management Plan are subject to endorsement by City Council. The main issue is whether a permit will be granted for raising the water levels by pump- ing from the Jock River at times of high flow. THE CANADIAN FIELD-NATURALIST Vol. 116 a Lac Leamy golf course: Letters were sent to the Ministers of Environment and Canadian Heritage. Our objections apply to the proposed construction of golfing greens on public land in forested areas, at minimal distances from the Lac Leamy wetlands bordering the Ottawa River, and the environmental impact of chemical run-off. Wetland drainage concerns: In essence, the problem is that existing regulatory controls tend to regard drainage as inherently beneficial, and do not require assessment of damage to significant wetlands that might be caused by drainage of adjoining land. Rosenbaum attended a work- shop held by the Ministry of Municipal Affairs and Housing on 4 October 2001 to review the Provincial Policy Statement (PPS) on land use. Main points made: (1) Clause 2.3.4 gives existing agriculture (even when recently-intro- duced) precedence over any consequences to natural land. (2) The guidelines apply to decision makers (planners and politicians) but not to developers and individuals if actions taken do not require a permit. Alfred Bog: Negotiations are apparently still ongoing, and no peat extraction is presently taking place within the official boundaries of the bog. Leitrim Wetlands: Philip Martin attended two meetings of the newly formed Leitrim Wetlands Advisory Committee, set up by South Nation Conservation Authority, and including repre- sentation from developers. We are unwilling to endorse any management plan until the future of the entire area of the pre- sent wetland has been resolved. Decisions by federal depart- ments (Fisheries and Environment) are delayed, perhaps due to new Transport Canada Risk Assessment investigations on the toxic dump, suggesting increased concern over possible liability if a subdivision is built. S. ROSENBAUM Education and Publicity Committee The committee organized presentations for the national wildlife festival, earth day, environment week, the OC ele- mentary teachers PE Day and Leamy Lake appreciation day. Eight nature walks were conducted and two slide presentations prepared for various groups, including the Trinidad and Tobago Field Naturalist Club, where Tom Bearss, the acting Canadian High Commissioner, spoke on our behalf. Members of the committee served as judges at the OC Regional science fair. Media relations students at Carleton University were assisted with a class project. Over $500 was raised through sales of club merchandise at outside events and club meetings. Publicity was provided to support club activities and volunteer recognition. P. PETERKIN Excursions and Lectures Committee Excursions and lectures organised monthly evening pre- sentations and field trips throughout the year. The two most significant events were the resignation of Ellaine Dickson (due to ill health) from the Committee after 33 years of service, and the cancellation of the bi-annual trip to Pt. Pelee. This was due to our local Leamington contact booking the wrong dates. The committee is now working on a plan that is 3-6 months ahead of schedule. R. JOHN os | 2002 MINUTES OF 123RD ANNUAL BUSINESS MEETING 517 Executive Committee The Executive Committee met once in 2001 to discuss how the OFNC should handle the request from the Canadian Nature Federation for help in organizing its 2002 annual general meeting — specifically, that OFNC would propose a roster of speakers and trip leaders to the CNF. Executive Committee decided to refer the matter to the Excursions and Lectures Committee, who will set up an ‘ad hoc’ committee to help the CNF as per their request. E. ZURBRIGG Finance Committee 1. The Committee met on three occasions.. In December 2000 the Committee examined the prelimi- nary (pre-audit) Financial Statement for 1999/2000. The audited Statement was approved at the Club’s January 2001 Annual Business Meeting. The Statement showed a signifi- cant loss on OFNC account offset by a substantial surplus on CFN account. At a meeting in April the Committee reviewed the half-yearly financial performance of the Club. In August, the Committee prepared a draft budget for 2001- 2002. The draft, which proposed a balance of revenue and expenditure for both the CFN and OFNC accounts, was presented to Council in September and adopted at its October Meeting. The Committee recommended no change in the membership fee structure. A. HALLIDAY Fletcher Wildlife Garden Committee The level of activity at the Fletcher Wildlife Garden remained steady in 2001, compared with the previous year. The Management Committee met monthly, and volunteers put in about 2000 hours of work. Members of the Friday morning Backyard Garden crew again gave unstintingly of their time every week from April to November to improve this showpiece of the garden. Weekend work days helped with general maintenance and the annual spring plant sale and exchange raised funds for garden operations. Our summer student proved to be capable, enthusiastic and hard-working. Much of her time was spent on invasive plant control, but she also helped build a fence, repainted our sign, carried out Internet searches and helped visitors, including a number of tourists who discovered the FWG after reading a write-up in Ottawa Magazine. The Interpretation Centre was open for several hours on week- days and was staffed on Sunday afternoons by volunteers. Exhibits inside were revised and enlarged and the floor was repainted. The centre was broken into twice, but with no significant loss of contents. A telephone (613-234-6767) was installed in the spring to provide a message system, emergency communication, and Internet access. Carleton University has continued with the swallowwort project, doing basic biology and testing non-chemical con- trol methods on test plots. Notices to explain the project were erected, although the material was vandalized repeat- edly. Wildlife gardening publications and newsletters were produced and disseminated to the public; these were funded by a grant obtained from the Canada Trust Friends of the Environment Program. The FWG continued to be a central location for OFNC activities, Council and at least six com- mittees and subcommittees meet regularly in the Interpretation Centre, as does the Ottawa Stewardship Council. Although the FWG’s financial position is quite good, all income in 2001 came from the OFNC allocation, the Taverner Cup, and donations from individuals. The committee would like to apply for funding for better sig- nage, but is constrained by the lack of a formal agreement with Agriculture and Agri-Food Canada. Although the FWG has a roster of enthusiastic volun- teers, the management committee continues to shrink, making it difficult to organize and supervise work parties. D. HOBDEN Macoun Field Club Committee The committee met five times to plan the weekly meet- ings for the children and young people of the Macoun Field Club through the course of the school year. Committee members arranged speakers and workshops for the Friday afternoon and Saturday morning meetings, and they led 16 field trips and 3 camping trips. Membership was a concern only in the two youngest age-groups, which had to be combined. There were 22 Juniors/Intermediates (8-13 years of age) and 20 Seniors (high school age) involved in the Macoun Club this year. R. LEE Membership Committee The distribution of memberships for 2001 is shown in the table (below), with the comparable numbers for 2000 in brackets. These statistics do not include three complimenta- ry memberships awarded to winners of the 2001 Science Fair competition nor the 23 affiliate organizations which receive copies of the Club’s publications. Distribution of Memberships in The Ottawa Field-Naturalists’ Club CANADIAN Type Local Other Family 345 (350) St (23) Individual 366 (340) bed (122) Honorary 17 (15) 8 (8) Life 21 (22) oe aa) Sustaining 8 (8) 2 (1) Total 797 C735) 184 (175) FOREIGN USA Other Total 32) 2 (1) 381 (376) 22 (25) 4 (4) 513 (491) 0 (0) 0 (0) 25 (23) 5 (5) CL) 49 (49) 0 (0) 0 (0) 10 (9) 30 (32) 7 (6) 978 (948) a ee ee Te 518 THE CANADIAN FIELD-NATURALIST This year the Club awarded Honorary Memberships to five persons. Three prominent and long-time members of the Club passed away this year. Dr. D. E. McAllister was an Honorary and member of the Club since 1958. Patricia (Mickey) Narraway was a member of Council for many years and a member of the Club since 1967. Dr. W. E. Ricker had been a member of the Club since 1930, perhaps the longest tenure in the history of the Club. D. SMYTHE Publications Committee The Publications Committee met twice in 2001. Three issues of The Canadian Field-Naturalist were pub- lished in 2001; Volume 114, #4, and Volume 115, #1, 2. These three issues contained in 598 pages: 46 articles; 23 notes; 4 COSEWIC articles; 28 book reviews; 147 new titles; 3 commemorative tributes; 12 pages of News and Comment; Vol. 116 and a 34-page index. An Associate Editor for mammalogy, W. Ballard, retired and was sent a certificate of appreciation for his seven years of service. There were no other changes in the panel of Associate Editors. Five articles received support from the Manning Memorial Fund, for a total of eight to date. Volume 35 of Trail & Landscape was published in four issues containing 228 pages. This was the largest volume in some years because it contained two lengthy articles: Consaul, Dugal, and Boles on the Leitrim wetlands and Coad on a key to the fishes of the National Capital Region; and a five-year index. Fenja Brodo retired as Editor and Sandra Gushue as Production Manager with the publication of Issue 1, both after ten years of service. They have remained on the staff as Associate Editor and Production Assistant, respectively. With Issue 2, Karen McLachlan-Hamilton took over as Editor. The Club owes a huge debt of gratitude to all of these people. R. BEDFORD 7 ] ‘ ) : 2002 Auditor’s Report MINUTES OF 123RD ANNUAL BUSINESS MEETING To The Members of THE OTTAWA FIELD-NATURALISTS’ CLUB I have audited the balance sheet of THE OTTAWA FIELD-NATURALISTS’ CLUB as at September 30, 2001, the statement of changes in net assets, and the state- ments of operations. These financial statements are the responsibility of the organization's manage-ment. My responsibility is to express an opinion on these statements based on my audit. Except as explained in the following paragraph, I conducted my audit in accordance with generally accepted auditing standards. Those standards require that I plan and perform an audit to obtain reasonable assurance whether the financial statements are free of material misstatement. An audit includes examining evidence supporting the amounts and disclosures in the financial statements. An audit also includes assess- ing the accounting principles used and significant esti- mates made by management, as well as evaluating the overall financial statement presentation. In common with many non-profit organizations, The Ottawa Field-Naturalists' Club derives some of its revenue from memberships, donations, and fund rais- ing activities. These revenues are not readily suscepti- ble to complete audit verification, and accordingly, my verification was limited to accounting for the amounts reflected in the records of the organization. In my opinion, except for the effect of the adjust- ments, if any, which I might have determined to be necessary had I been able to satisfy myself concerning the completeness of the revenues referred to in the preceding paragraph, these financial statements pre- sent fairly, in all material respects, the financial posi- tion of the OFNC as at September 30, 2001, and the results of its operations and changes in net assets for the year then ended in accordance with generally accepted accounting principles. JANET M. GEHR Chartered Accountant North Gower, Ontario December 18, 2001 The Ottawa Field-Naturalists’ Club Balance Sheet September 30, 2001 ASSETS CURRENT (Sasi (NOLS TR ee ante ws Investment certificates (Note 1)... Marketable securities (Note 2)..... ACCOUNTS TECEIVADIE ..00..6.csc-ccesee: PREP AIG CAPEMSES 0520 cadens CAPITAL ASSETS (Note 3) Land — Alfred Bog.................... LIABILITIES AND FUND BALANCES CURRENT Accounts payable and BECIUE IMAPUITIES ..-: caseserce concen Sterne dG VENUS .cicesorsceaveereeeeees [:ife micmibersiips 5 .....00...e0e--caane NET ASSETS WWinnesiticte dat tres 54sec eee (CUM RESERVE oto soicesee odie cic dan-tceeske Manning, primcipall........::........--.. Manning interest - OFNC — CFN aye) 0 (2) 01 0 2] a Pe ee Ane Anne Hanes Memorial................. de Kiriline=awrenCe ......cts0c...0ces Macoun Baillie Birdathon ........... PICS assncatantcanapetiuestasins 2001 $ 50,760 110,219 194,234 5,386 3,348 $ 364,947 $ 2,000 13,788 15,788 he Wr pe | 99,427 100,000 100,000 2,186 8,225 1,569 870 24,553 (7) 615 337,438 $ 364,947 a19 2000 $ 9,519 106,939 185,765 19,689 1,000 522.012 3,348 $326,260 $ 2,000 13,532 15.332 11,249 66,721 100,000 100,000 1,203 4,813 1,308 870 22,941 1,039 584 299,479 326,260 TTT Egy ahem nak pee ee Smee rian a 520 THE CANADIAN FIELD-NATURALIST Vol. 116 The Ottawa Field-Naturalists’ Club The Ottawa Field-Naturalists’ Club Statement of Operations Statement of Operations — For the Year Ended September 30, 2001 The Canadian Field-Naturalist 2001 2000 For the Year Ended September 30, 2001 2001 2000 REVENUE Mismibersbigs: ..28ei2 2c..<. 7 a Pures eMC TERS oo oc cdceensteenneee se 455 1,309 97,898 97,498 iG) C175) 0] Lt ge RR ae ps RL a 534 444 IVICHIIERS IED oo. chee cee e, ease cg 1,365 L732, EXPENSES CRIES ASSISIAIG ons cac, akachcccsscre 1,000 1,000 Publishiniow.s. s:8..62 eee 35,108 58,461 HV ENS7|110)v eget a 2 1,593 1,206 Reprintsst educetecele J yie gs Ae 9,430 6,135 | TPT er ey eae ROLE Re eae 688 655 Cine ml atte: en Pern. Bess thee 7,942 6,070 Fe TUT eet elt. SE 1,000 1,000 [SCT 60 Ce Ome, Ree e i CTE 3,628 3,258 Co | (APN Se Bee av alice: Eero 1505 1,494 Olfice ASSISEANE: goose 5,000 5,000 4) F675) 0211 Te ar OR 782 230377 FIQMOANIA e555 oat ac ceeeeaes 6,000 6,000 (0 apiAlaw hares “vel th tds Mrysinn A * ob 3,867 6,045 8922, _ 10877 _ Meier Ee Pa i ies 1,028 2,020 CLUB ACTIVITY EXPENSES 72,003 92,989 PC Css eee LAE SA ee 280 DES Sr eg eee ne 466 875 EXCESS REVENUE OVER Education and Publicity .............. 182 387 EXPENSES _ $25,895 _ $4,509 _ Excursions and Lectures............... (311) (98) Macoun Field:Clup ........0....cccecc.0. 955 List "5 2 yee oR te OR 65 429 Drath de (eandscape «..ssccccatvscns Teas *- - — ~ ih. »” : ate A roe “=a a 4 ts ee ES 8 ag OP “= - Volume 116, Number 4 October—December 2002 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Her Excellency The Right Honourable Adrienne Clencien: CC. CAME, © ine Governor General of Canada His Excellency John Ralston Saul, C.C. 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 infor- mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain- ing or restoring environments of high quality for living things. Honorary Members Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope Donald M. Britton R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. Irwin M. Brodo Anthony J. Erskine | Hue N. MacKenzie Joyce and Allan Reddoch William J. Cody John M. Gillett Theodore Mosquin Mary E. Stuart Francis R. Cook C. Stuart Houston Eugene G. Munroe John B. Theberge Ellaine Dickson George F. Ledingham Robert W. Nero Sheila Thomson 2002 Council President: Eleanor Zurbrigg Vice-President: Gary McNult Ronald E. Bedford Francis R. Cook David W. Moore a4 eee : Rosanne Bishop Barbara Gaertner Robert Roach : Irwin Brodo Diane Lepage Stanley Rosenbaum Recording Secretary: Ken Allison John Cameron Diane Holms Louise Schwartz Treasurer: Frank Pope William J. Cody David Hobden David Smythe Kathy Conlan Beverly McBride Dorothy Whyte To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, Box P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net. For information on Club activities telephone (613) 722-3050 or check http//www.achilles.net/ofnc/index.htm The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. We acknowledge the financial support of the Government of Canada toward our mailing cost through the Publication Assistance Program (PAP), Heritage number 09477. Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1RO; (613) 269-3211; e-mail: feook @achilles.net Copy Editor: Elizabeth Morton Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 759-1374 Book Review Editor: Dr. J. Wilson Eedy, 9074-32 Side Road, R.R.1, Glencairn, Ontario LOM KOC Canada e-mail: terfa@ geoniger.com Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen Charles D. Bird Brian W. Coad Donald F. McAlpine Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr. Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication should be addressed to the Editor and sent by postal mail. Exception: book reviews should go directly to Book Review Editor. Subscriptions and Membership Subscription rates for individuals are $28 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $28 (individual) $30 (family) $50 — (sustaining) and $500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and mem- — bers (including USA) must add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers — the Ottawa District and Local Club events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at $28 per year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate P.O, Ottawa, Canada K1Z 1A2. Canada Post Publications Mail Agreement number 40012317. Return Postage Guaranteed. Date of this issue: October-December 2002. (May 2003). Cover: Brittle Prickly-pear Cactus, Opuntia fragilis, at Mellon Creek, Kaladar, Ontario, (SJD 50/4 DAO) photographed 27 June 1999 by Stephen J. Darbyshire. See Staniforth, Cody, and Frego pages 4547-550. SMITHSOA IN an Field-Naturalist The Canac Volume 116, Number 4 At hi Fas eS a a Te * U October—December 2002 Rapid Estimation of Plant Biomass Used as Forage or Cover by White-tailed Deer, Odocoileus virginianus, and Snowshoe Hare, Lepus americanus, in Mixed and Coniferous Forests of Southeastern Québec ISABELLE ROULEAU!, MICHEL CRETE!.23 GAETAN DAIGLE4, PIERRE ETCHEVERRY!, and CHANTAL BEAUDOIN? 'Département de biologie, Université du Québec a Rimouski, 300 Allée des Ursulines, Rimouski, QC, GSL 3A1 Canada 2Société de la faune et des parcs du Québec, 675 boul. René-Lévesque Est (BP 92), Québec, QC, GIR 5V7 Canada 3Département de biologie, Université Laval, Sainte-Foy, QC, G1K 7P4 Canada 4Département de mathématiques et de statistique, Université Laval, Sainte-Foy, QC, GIK 7P4 Canada Rouleau, Isabelle, Michel Créte, Gaétan Daigle, Pierre Etcheverry, and Chantal Beaudoin. 2002. Rapid estimation of plant biomass used as forage or cover by White-tailed Deer, Odocoileus virginianus, and Snowshoe Hare, Lepus ameri- canus, in mixed and coniferous forests of southeastern Québec. Canadian Field-Naturalist 116(4): 523-528. Estimation of vegetation biomass in forest ecosystems is time-consuming because of inherent high variability. We devel- oped 10 linear regression models to predict standing dry biomass of most plant taxa potentially used as forage or cover by White-tailed Deer and Snowshoe Hare in forests of southeastern Québec. Vertical interception and lateral coverage, which served as explanatory variables in regressions, were measured along 2-m transects before clipping, drying and weighing vegetation. Plant species with similar morphology and size were pooled. Although we collected vegetation samples from two areas 400 km apart, we could combine data for regressions except for low herbs (< 30 cm). Models covered a wide range of biomass and fitted the data reasonably well when In-transformed, with R? values varying between 0.61 and 0.85. Cross-validation confirmed model robustness. Key Words: biomass, boreal, cover, forage, forest, linear regression, method, mixed, Québec, technique L’estimation de la biomasse végétale dans les écosystémes forestiers demeure laborieuse a cause de son inhérente grande variabilité. Nous avons développé 10 modéles de régression qui prédisent la biomasse de la plupart des taxa végétaux poten- tiellement utilisés comme nourriture ou couvert par le cerf de Virginie et le li¢vre d’ Amérique dans les foréts du sud-est québécois. L’interception verticale et le recouvrement latéral des espéces végétales, qui ont servi de variables explicatives dans les modéles, furent mesurés au-dessus de parcelles linéaires de 2 m, avant qu’on coupe, séche et pése les végétaux. Les espéces végétales avec un port et une taille semblables ont été regroupées. Bien que les échantillons fussent recueillis dans deux régions séparées de 400 km, il fut possible de les regrouper sauf pour les herbacées basses (< 30 cm). Les modéles ont couvert une gamme étendue de biomasse et se sont ajustés relativement bien aux données aprés une transformation logarith- mique, avec des R? compris entre 0,61 et 0,85. Une validation croisée a confirmé la robustesse des modéles. Mots clefs: biomasse, boréale, faune, forét, mélangée, méthode, Québec, régression linéaire, technique. During concurrent studies on the summer habitat of White-tailed Deer (Odocoileus virginianus; Rouleau et al. 2002) and the space used by Snowshoe Hare (Lepus americanus) in summer and in winter (Beaudoin 2001), we were confronted by the need to quantify forage and cover availability. We wanted to estimate plant biomass because direct measurements of vegetation represent a prerequisite for improving our understanding of wildlife-habitat relationships (Morrison et al. 1998: 151). The large spatial variabil- ity of plant biomass (e.g., Créte and Jordan 1982; Créte 1989 ; Créte and Courtois 1997) requires large sample sizes, which are time-consuming. As shoulder height of adult White-tailed Deer averages 92 cm (Banfield 1977), 1.5-m high vegeta- tion suffices to completely hide this species. When foraging, White-tailed Deer zigzag slowly and rarely browse plants above 1.5 m from the ground. Snowshoe Hare can feed on twigs up to 45 cm above snow surface when standing on their hind feet (Banfield 1977). When resting or standing on their feet, hare require dense vegetation up to around 30 cm to be hidden from the view of terrestrial preda- tors. Hare are well adapted for locomotion in the snow (Murray and Boutin 1991), and generally feed and move on the top of the snow pack. Snow depth WwW 524 varies between 50 and 100 cm during winter in southeastern Québec (e.g., Dumont et al. 2000). When stratifying the vegetation likely to serve as food or cover for White-tailed Deer and hare, we then used three practical thresholds for height class- es: 30 cm, 50 cm, and 150 cm. Methods exist to estimate vegetation volume or biomass quickly with auxiliary variables which can be measured rapidly (e.g., Bonham 1989; Créte et al. 1990b), but none were adapted to our needs. In this study, we wanted to determine whether we could pre- dict, with simple and rapid measurements, plant biomass growing in the 0-1.5 m stratum for mixed and coniferous forests of varying ages in south-eastern Québec. As we tallied 214 taxa of herbs and woody species in the deer range in Québec (Boucher 2003), we had to pool species. based on their growth form to obtain adequate sample sizes for each regression model. Study Area and Field Methods We studied plant biomass in the Bas-Saint- Laurent (~48° 00’ N; ~69° 00’ W) and Montérégie (=46° 00’ N; =73° 00’ W) regions of Québec during the growing seasons of 1998 and 1999. The Bas- Saint-Laurent region occupies a transition zone between the northern hardwood forest and the boreal forest; Sugar Maple (Acer saccharum) and Yellow Birch (Betula alleghaniensis) often grow on hilltops and well-drained hillsides, whereas Balsam Fir (Abies balsamea), White and Black spruce (Picea glauca, P. mariana) frequently dominate with Trembling Aspen (Populus tremoloides) and White Birch (Betula papyrifera) elsewhere (Rowe 1972). Most forest stands have been affected by logging, insect epidemics, or fire during the last century. The milder climate in Montérégie favours the growth of a greater variety of tree species, including Sugar Maple, Yellow Birch, White Ash (Fraxinus ameri- cana), American Linden (Tilia americana), Ironwood (Ostrya virginiana), White Pine (Pinus strobus) and Hemlock (Tsuga canadensis; Rowe 1972). Commercial logging appears to have been less important in Montérégie than in Bas-Saint- Laurent, likely due to a fragmented pattern of land ownership, but human activity has affected most for- est stands. Aerial surveys carried out in early winter revealed that White-tailed Deer density averaged 1 individual km in Bas-Saint-Laurent (1999) and 12 individuals km? in Montérégie (1997). Deer brows- ing likely reduced plant biomass in the latter region, while browsing impact remained negligible in the former (Rouleau et al. 2002). We collected vegetation samples during the grow- ing season between 3 June 1998 and 31 August 1999. Green-up begins in early May in Montérégie and two or three weeks later in Bas-Saint-Laurent (Lesage et al. 2000). Samples came from sites used THE CANADIAN FIELD-NATURALIST Vol. 116 - by radio-collared White-tailed Deer (n = 109) and Snowshoe Hare (n = 23), or from 17 grids used to estimate small mammal and hare density (n = 66). Seven observers participated in the study. We used a line-intercept technique to estimate veg- etation biomass (e.g., Créte et al. 1990a). Sampling plots were 2 m long and the location of their starting point coincided with a previous telemetry location of White-tailed Deer or hare, or a predetermined point on permanent grids used for estimating hare or small mammal density. Transect orientation was selected randomly. Observers located sampling plots with a hip chain and a compass. We made a length and height assessment of plant coverage by species (genus or family, depending on ease of taxonomic identification). To facilitate data collection, plants up to 150 cm tall were recorded in two sequences. First, we recorded lycopods, trees and shrubs < 50 cm tall and all herbs. Secondly, we recorded all the foliage from 0-150 cm on trees and shrubs that exceeded 50 cm in height. We omitted lichens and mosses due to their rarity in both regions. We placed a 1-1.5 cm wide measuring tape on the ground surface for recording measurements, using the metric side as the sampling line. The observer stood perpendicular to the sampling line and record- ed the position of the beginning and end of each vertical interception of the line by species, genus, or family to the nearest cm. The difference between the end and beginning of each interception yielded the 1.5m 0.5m 1m 2m Lateral view FiGuRE 1. Schematic representation of vertical interception and lateral coverage measurements taken on three shrubs with foliage intercepting the sampling line (dotted line on vertical view) and surface area (ver- tical view) from which foliage was clipped, dried and weighed (0.1 m X 2 m). See text for details. 2002 length (L; Figure 1), with a minimum of | cm (i.e., if L < 1, then L = 1). The observer then estimated, from a lateral view, the average height (H) of the green plant material associated with each intercep- tion within a 5-cm strip on each side of the sampling line (see below). H included leaves and stems of herbs but only foliage of woody species (Figure 1). We did not distinguish individual plants for herba- ceous or woody species, so a single plant could pro- vide several measurements if line interceptions origi- nating from the same stem were separated by an unoccupied space (Figure 1). We then collected the green vegetation present within 5 cm of both sides of the sampling line (0.2 m7) and from the ground to a height of 150 cm (0.3 m*). We used wooden poles and elastic cords to delimit the sampling volume. We harvested herbs with pruning shears from the ground when originat- ing in the sampling surface. We collected only the leaves of deciduous woody species whereas we col- ROULEAU, CRETE, DAIGLE, ETCHEVERRY, AND BEAUDOIN: PLANT BIOMASS 525 lected needle-bearing twigs plus needles for conifers. Samples were sorted by species and placed in plastic or paper bags for transportation, except for woody deciduous species > 50 cm, whose leaves were pooled. Vegetation samples were dried in an oven at 40-60°C until weight stabilisation and weighed with a precision of + 0.01 g. Statistical Analysis To obtain adequate sample sizes, we pooled data into 10 categories based on morphological and size similarities (Table 1); we used species description by Marie-Victorin (1995) to partition species into cate- gories. We In-transformed all biomass data because distributions were log-normal. The validity of this transformation was tested by fitting a generalised gamma model to the data and testing at a = 0.05 for the shape of the parameter to be equal to 0 with the Lagrande multiplier chi-square statistic (Proc Lifereg, SAS 1989). Furthermore, this trans- TABLE 1. Species composition of 10 plant categories with morphological similarities and likely to serve as forage and cover for wildlife in mixed and coniferous forests of northeast North America. Plant categories Herbs < 30 cm (Bas Saint-Laurent) Species composition Clintonia borealis, Coptis groenlandica, Chiogenes hispidula, Cornus canadensis, Epigaea repens, Fragaria sp., Hieracium sp., Linnaea borealis, Maianthemum canadense, Oxalis montana, Pyrola sp., Taraxacum sp., Trientalis borealis, Viola sp. Herbs < 30 cm (Montérégie) Circea lutetiana, Clintonia borealis, Coptis groenlandica, Cornus canadensis, Dalibarda repens, Fragaria sp., Maianthemum canadense, Mitchella repens, Mitella diphylla, M. nuda, Oxalis montana, O. stricta, Taraxacum sp.., Tiarella cordiafolia, Trientalis borealis, Viola sp. Herbs = 30 cm Anaphalis margaritacea, Aralia nudicaulis, A. hispida, Aster sp., Centaurea nigra, Epilobium angustifolium, E. glandulosum, Eupatorium rugosum, Impatiens capensis, I. sp., Lycopus americanus, Lysimachia nummularia, L. terrestris, Mimulus ringens, Ranunculus acris, Senecio sp. Solidago sp., Thalictrum sp. Ferns Athyrium filix-femina, Dennstaedtia punctilobula, Dryopteris sp., Matteuccia struthiopteris, Onoclea sensibilis, Osmunda cinnamomea, O. claytoniana, O. regalis, Pteridium aquilinum Graminoids Deciduous seedlings (< 50 cm) Cultivated and native Graminae, Cyperacae Acer sp., Alnus rugosa, Amelanchier sp., Betula papyrifera, B. populifolia, B. sp., Corylus cornuta, Fraxinus americana, Ostrya virginiana, Populus tremu- loides, Prunus serotina, Tilia americana, Ulmus americana Small deciduous shrubs (< 50 cm) Cornus stolonifera‘, Diervilla lonicera, Kalmia angustifolia, Lonicera canadensis“, Myrica gale’, Nemopanthus mucronatus“, Ribes sp., Rhododendron canadense, Rhus radicans, Rubus sp.*, Spiraea latifolia’, Vaccinium sp. Large shrubs and trees (deciduous; > 50 cm) Acer pensylvanicum, A. rubrum, A. saccharum, A. spicatum, Alnus rugosa, Amelanchier sp., Betula populifolia, Corylus cornuta, Fraxinus americana, Ledum groenlandicum, Populus tremuloides, Prunus pensylvanica, P. serotina, Rhamnus alnifolius, Viburnum alnifolium Balsam Fir Abies balsamea Spruces White Cedar Thuja occidentalis Picea glauca, P. mariana, P. rubens “Some samples were included in the categorie “Large shrubs and trees” when >50 cm : 526 THE CANADIAN FIELD-NATURALIST formation generally minimised the error sum of squares among all transformations in the Box-Cox family. We used two independent variables to predict plant biomass per sampling plot: L and H. L represented the sum of the interception lengths per plot for a plant species, genus, or family (i.e., XL,: Figure 1), while H was the mean weighted height (i.e., 2» H,*L,/L: Figure 1). We considered all combinations of the two independent variables (non- transformed, In(X), X?, X'”, etc.), including polyno- mial relationships, to select the regression models that best fitted the biomass data. We chose as the final model the most parsimonious one, i.e., the model with the highest adjusted R* which included only significant explanatory variables (P < 0.05), ensuring that residuals were normally distributed (Shapiro-Wilk test), homogenous and without pat- terns (visual examination of the plot). In addition, we used the Dffits, Hatdiag and CovRatio statistics (Proc Reg, SAS 1989) to detect influential data points. We introduced indicator variables (Neter et al. 1985) and interaction terms into the regression models to test whether we could pool observations from the two study regions. We used leave-one-out cross-validation (Stone 1974) to check the robust- ness of regression models. We systematically excluded one data point at a time and predicted its value with the regression model adjusted with the remaining data set (SAS 1989). We then computed Vol. 116 - the coefficient of determination between observed and predicted values to ascertain how they matched. Results Apart from lycopods, for which we could not fit a model due to the small sample size (n = 7), we found highly significant (P < 0.0001) regression models predicting dry plant biomass for all plant categories (Table 2). We collected samples in both regions for all plant categories except coniferous trees (fir, spruces and White Cedar: Thuja occidentalis). Our statistical analysis revealed that we could pool sam- ples from the two regions, except for low herbs, for which the intercept differed (P < 0.0001: Table 2). The regression models explained between 61 and 85% of the variability in biomass for the 10 plant categories. Only total length of vertical interception (L) significantly predicted biomass of low herbs, graminoids, deciduous seedlings and small decidu- ous shrubs, while both length of vertical interception and height of lateral coverage (H) were significant factors for ferns, tall herbs and large woody plants (Table 2). For Balsam Fir and White Cedar, parame- ter estimates for the two independent variables (L and H) did not differ (P > 0.05) so we used the prod- uct (LXH) to predict plant biomass. R? computed during cross-validation were generally 2-3% lower than those for models fitted with full data sets. TABLE 2. Characteristics of regression models developed to predict green biomass for 10 plant categories for the 0-150 cm height stratum in mixed and coniferous forests in 0.2 m? or 0.3 m? depending on whether expressed as surface southeastern Québec. Models predict dry mass in g per area or volume. The value, MSE/2, must be added to the predicted In biomass before transforming back to the normal scale to prevent bias (Ung and Végiard 1988). Plant group Regression model? Adjusted Mean Squared N Range in In (biomass) = R? (Cross- Error (MSE) biomass (g)° validation R?) Herbs < 30 cm (Bas-Saint-Laurent) = - 2.7647 + 1.0173 In (L) O.7Z(071) 0.6480 84 0.01-9.99 Herbs < 30 cm (Montérégie) - 4.0254 + 1.0173 In (L) 0.72 (0.71) 0.6480 58 = 0.01-1.09 Herbs = 30 cm - 2.7485 + 0.7296 In (L) 0.66 (0.63) 0.8451 81 0.01-57.93 + 0.3861 In(H) Ferns - 3.9042 + 0.9485 In (L) 0.75 (0.74) 0.7147 119 0.01-39.44 + 0.4110 In(H) Graminoids - 3.8217 + 1.3297 In (L) 0.79 (0.77) 0.7906 38 ~=—-0.01-30.75 Deciduous seedlings (< 50 cm) - 3.3175 + 0.9911 In (L) 0.61 (0.59) 0.9346 55. O0LTSs Small deciduous shrubs (< 50 cm) - 2.9772 + 0.9643 In (L) 0.65 (0.63) 0.6631 74 0.02-23.46 Large shrubs and trees - 3.5029 + 0.8960 In (L) 0.82 (0.81) 0.4159 49 0.05-44.64 (deciduous; > 50cm) + 0.4218 In (H) Balsam Fir - 3.0136 + 0.7503 In (L(H) (0.79 (0.77) 0.4798 46 ~=0.33-141.00 Spruces - 2.2202 + 1.0608 In (L) 0.81 (0.78) 0.3456 32 | 1L13-13%e7 + 0.2533 In (H) White Cedar - 3.7381 + 0.8008 In (LXH) 0.85 (0.82) 0.2894 25 0.66-82.48 *L = total length (cm); H = mean weighted height (cm); > Minimum and maximum values that were used to fit the models 2002 Discussion Modelling generally necessitates simplification of study systems and our work was no exception. We hypothesized that we could describe most variation in dry plant biomass from rapid measurements taken in two dimensions: vertical interception for length and lateral coverage for height (L and H, respective- ly). In addition, we assumed that biomass density and spatial arrangement remained relatively similar for plant taxa pooled within categories (Table 1). We conclude that such assumptions were valid, given that most adjusted R* values exceeded 0.70. Cross- validation confirmed model robustness. In general, biomass of low vegetation could be estimated using only vertical interception whereas biomass predic- tion of taller plants necessitated both vertical and lat- eral measurements. This may partly depend on the precision of our measurements (1 cm), which were proportionally more imprecise for small plants. Height also served to predict fern biomass probably because the species sampled (Table 1) exhibit large size variation. The more complex models for tall plants could also depend on more complex architec- ture than that of low vegetation. In general, biomass of conifers exceeded that of woody deciduous species by a factor of 2 to 10 for comparable L and H measurements. This result is likely due to the inclusion of twigs bearing needles for conifers in addition to the higher specific leaf area in coniferous than deciduous species (Lei and Lechowicz 1990; Sprugel et al. 1996; Reich et al. 1998). We only found one significant (P < 0.0001) regional difference when adjusting our models to predict plant biomass, and that was for herbaceous species smaller than 30 cm in height. The two regional models differed only in intercept, and pro- duced biomass predictions differing by a factor of 3.53. We believe that such a difference arose from different species composition (Table 1) and domi- nance, and from the high White-tailed Deer density in Montérégie, which has likely reduced plant size (e.g., Balgooyen and Waller 1995). The models that we computed are relatively robust because we used data collected by different crews in two different areas separated by approximately 400 km. Likewise, our models should produce reasonable estimates of biomass for most plant taxa growing in Ontario, Québec and in adjacent parts of southwest New York, Vermont, Maine, and New Brunswick. Further away, researchers or managers could use double sampling (Bonham 1988: 202) to alleviate potential biases. Our model can serve for studies on White-tailed Deer or Snowshoe Hare, but also for other animal taxa which use the 0-1.5-m high stra- tum in forest ecosystems. Users should consider White-tailed Deer density when predicting biomass of low herbs; we suggest using the model computed ROULEAU, CRETE, DAIGLE, ETCHEVERRY, AND BEAUDOIN: PLANT BIOMASS 527 in Montérégie if deer density exceeds 10 animals km. Although data collection is rapid, the use of our regression models necessitates careful field mea- surements to produce unbiased estimates. One should always imagine the volume of reference; i.e., 5 cm on each side of the transect line, when estimat- ing lateral coverage. In addition, users must tally only plant specimens with vertical interception; plants present in the sampling volume but not cross- ing the transect line must be ignored. We already successfully used our models for detecting habitat preferences in Snowshoe Hare (Beaudoin 2001) and White-tailed Deer (Rouleau et al. 2002.). Given that we pooled plant species to compute our models, they can serve to predict biomass of any species; e.g., of preferred forage species only. We did not measure the time necessary to sample plots, but it took approximately 5 to 20 min to survey a plot, depending on vegetation density. Our method, coupled with plotless tree measurements (e.g., Grosenbaugh 1952) and sapling counts in circu- lar plots, will allow a rapid and complete characteri- sation of forest stands (e.g., Beaudoin 2001). Our results illustrate that relationships between vegetation coverage and biomass are curvilinear (all biomass data In-transformed), and thus the use of linear indices of vegetation abundance may preclude find- ing significant wildlife-habitat relationships. Acknowledgements We express our gratitude to C. Bélanger, L. Bélanger, R. Dupin, M. Fournier, M. Lavoie, K. Lévesque, C. Rondeau and L. Senneville for taking measurements in the field and processing plant sam- ples. S. Cdté, C. Daigle, A. Dicaire, L. Lapointe and J.- P. Ouellet kindly commented on a previous draft of this manuscript. This study received financial support from the Société de la faune et des parcs du Québec, the Ministére des Ressources naturelles du Québec, the Fonds FCAR and from NSERC. I. Rouleau benefited from a Fonds FCAR scholarship during the study. Literature Cited Balgooyen, C. P., and D. M. Waller. 1995. 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Problémes d’inférence Statistique reliés a la transformation logarithmique en régression. Canadian Journal of Forest Research 18: 730-738. Received 1 December 2000 Accepted 18 January 2003 Selection, Size, and Use of Home Range of the Appalachian Cottontail, Sylvilagus obscurus Monica A. STEVENS! and RONALD E. BARRY Department of Biology, Frostburg State University, Frostburg, Maryland 21532 USA 'Present address: Department of Zoology, University of New Hampshire, Durham, New Hampshire 03824 USA Stevens, Monica A., and Ronald E. Barry. 2002. Selection, size, and use of home ce of the Appalachian Cottontail, Sylvilagus obscurus. Canadian Field-Naturalist 116(4): 529-535. Little is known about the ecology of the Appalachian Cottontail (Sylvilagus obscurus) in the northern portion of its range. We studied selection, size and use of home range for eight radio-collared Appalachian cottontails in western Maryland from February to July 1995. Home ranges were large and highly variable. For both sexes combined, median 100% mini- mum convex polygon (MCP), 95% MCP, and 95% adaptive kernel home ranges measured 5.4, 4.3, and 4.7 ha, respective- ly. Home ranges had greater amounts of understory cover and lesser slopes (<25°) than the surrounding landscape. Rabbits used portions of home ranges in close proximity (<2 m) to concealment cover, with canopy closure <95%, and with Blackberry, Rubus alleganiensis, as the dominant shrub species; they avoided places >10 m from concealment cover, with low stem density (<20 stems/l-m radius plot), and with 95-100% canopy closure. Dense understory concealment and escape cover seemed to be more important than species of vegetation to Appalachian Cottontails in Maryland. Our results suggest that fragmentation of habitat could limit the distribution of the species. We also demonstrate the importance of analyzing individual variation of life history characteristics in addition to measures of central tendency. Key Words: Appalachian Cottontail, Sylvilagus obscurus, cover, habitat, home range, individual variation, adaptive kernel, minimum convex polygon, radiotelemetry. The Appalachian Cottontail (Sylvilagus obscurus) occurs from northeastern Pennsylvania, south through the Appalachian Mountains to northern Georgia (Chapman et al. 1992). During the past half century, however, the numbers and ranges of this and its more northern sister species, S. transitionalis (New England Cottontail), have decreased (Merritt 1987). Sylvilagus transitionalis [which some believe is conspecific with S$. obscurus] is considered nation- ally “vulnerable” (IUCN 2002*), and S. obscurus is “highly rare ... critically imperiled” in Maryland (Maryland Wildlife and Heritage Division 1994%*). The decline of both species is thought to reflect (1) gradual climatic changes since the last glaciation, (2) the subsequent reinvasion of lowland areas by the more ubiquitous Eastern Cottontail (Sylvilagus flori- danus) (Chapman et al. 1992), and (3) destruction, fragmentation, and maturation of habitat (Barbour and Litvaitis 1993; Probert and Litvaitis 1996). Few investigators (Chapman and Paradiso 1972; Chapman and Morgan 1973; Chapman et al. 1977; Chapman and Stauffer 1981; Spencer 1985) have studied the ecology of the Appalachian Cottontail in Maryland and West Virginia. No study has defined home ranges of S. obscurus in any part of its range. Home ranges of S. transitionalis in Connecticut estimated using live trapping and mark-recapture measured 0.2-0.7 ha (Dalke 1937). By contrast, home ranges of S. floridanus determined using radioteleme- try range from 1.0—2.8 ha (Chapman et al. 1982; Dixon et al. 1981), and males generally have larger home ranges than females (Trent and Rongstad 1974). * See Documents Cited section 5 Habitat of Appalachian Cottontails in southeast Virginia has been described as six- to seven- year- old clear cuts (Blymer 1976). In Maryland, however, this species is reportedly limited to high elevations (above 570 m) in habitats containing conifers and ericaceous (heath-like) vegetation (Chapman and Paradiso 1972; Chapman and Stauffer 1981). Barbour and Litvaitis (1973) found New England Cottontails in New Hampshire by searching sites with dense understory cover. Habitat fragmentation and patch size are important considerations in habitat requirements. S. obscurus in western Maryland may not survive and reproduce in forested areas less than about 20 000 ha (Chap- man et al. 1977). New England Cottontails on small patches (< 2.5 ha) had lower survival rates than rab- bits on large patches (>5.0 ha) owing to quality of habitat and forage (Barbour and Litvaitis 1993; Villafuerte et al. 1997). On a larger scale, New England Cottontails had lower survival rates in frag- mented landscapes than in more even landscapes (Brown and Litvaitis 1995). The objectives of our study were to determine size and habitat characteristics of home-ranges of Appalachian Cottontails in western Maryland. We tested the following hypotheses: (1) home ranges are similar in size (~0.2-0.7 ha) to those reported previ- ously for S. transitionalis; (2) males have larger home ranges than females; (3) rabbits select home ranges with extensive coniferous and ericaceous veg- etation; and (4) rabbits use microhabitats within the home range with the greatest amount of understory (concealment) cover. 29 530 THE CANADIAN FIELD-NATURALIST Study Area and Methods Study sites Our study was conducted in the Savage River State Forest, Garrett Co., Maryland. The forest encompass- es about 21 400 ha of woodlands and is one of the largest blocks of forest remaining in Maryland. It is primarily an even-aged, second-growth forest, about 70 to 90 years old. Sylvilagus obscurus is considered rare in the forest (Maryland Department of Natural Resources 1992*). Study sites were chosen on the basis of accessibility, reports of historical inhabitance by S. obscurus (Chapman and Morgan 1973; R. P. Morgan, personal communication; and S. A. Olcott, personal communication), the presence of habitat broadly characteristic of habitat used by S. obscurus (e.g., high elevations and abundant ericaceous vege- tation), lack of open corridors through which S. flori- danus could easily invade, and the presence of rabbit tracks and pellets. Our study site was located north of Turkey Lodge Ridge, 3 km east of New Germany State Park (79°05'W, 39°37'N). Elevation ranged from 730-820 m. The site was classified as predominantly mature mixed-oak forest; however, abundant patches of nine-year-old clear cuts existed (J. Hare personal communication). Mature stands were dominated by Chestnut, White, and Red oaks (Quercus prinus, Q. alba, and Quercus spp.), Red Maple (Acer rubrum), and Black Birch (Betula lenta), with an understory dominated by Mountain Laurel (Kalmia latifolia) and blueberries (Vaccinium spp.). Early successional stages after clearcuts were dominated by Mountain Laurel, blueberries, and Blackberry (Rubus alle- ganiensis). Conifers present were Eastern Hemlock (Tsuga canadensis) and White Pine (Pinus strobus). Procedure Rabbits were captured with wooden box traps (18 x 22 X 60cm) baited with apple slices. Traps were placed in areas where rabbit tracks and pellets were found. Trapping was performed from December 1994 to mid-March 1995. We distinguished S. obscurus from S. floridanus using morphological characteristics designed originally to distinguish S. transitionalis from the latter (Litvaitis et al. 1991). Individuals were anesthetized by intramuscular injection of ketamine hydrochloride (30-45 mg/kg body mass) and then ear-tagged (National Band and Tag Co., Newport, Kentucky). Blood samples were taken from the ear, centrifuged to isolate plasma, and stored at -80° C. Positive species identifications were obtained by electrophoretic analysis/isoelectric focusing of blood serum proteins (Chapman and Morgan 1973; Sommer 1997) using Ampholine PAGplates (Pharmacia LKB Biotecknology, Uppsala, Sweden) with pH gradients of 5.5 to 8.5. Adults (>750 g for females and >800 g for males: Chapman and Morgan 1973) were fitted with collar- mounted transmitters (Lotek Engineering, Inc., Vol. 116 - Newmarket, Ontario, Canada). We tracked radio- collared individuals using a hand-held H-antenna (Lotek Engineering, Inc., Newmarket, Ontario). Rabbits were approached on foot. Precise locations were confirmed most often by direct observation of the subject, or by carefully monitoring the strength of the radio signal to determine the cluster of vegeta- tion in which the subject was located. Rabbits initial- ly remain motionless when threatened (Whitaker 1980) so rabbit locations did not likely represent locations to which they fled due to our approach. We obtained an average of two locations per week, for durations of 2.5—5 months for individual rabbits. Rabbits were monitored between February and July 1995, i.e., during the breeding season, which extends from February to September. Days were divided into six 4-hour blocks, and equal num- bers of locations were obtained for each time block that contained daylight hours. Universal-Transverse-Mercator-grid coordinates used to calculate home-range sizes were determined for each location using GPS (Trimble Navigation Plus, Trimble Navigation, Ltd., Sunnyvale, California). A total of 180 readings was taken at each location and then averaged using the software PATHFINDER (Trimble Navigation, Ltd., Sunnyvale, California). Coordinates had mean error polygons of 0.34 ha, as determined by obtaining 35 sets of GPS coordinates for seven known points. This error was consistent with errors in home-range estimates for mammals with home-range sizes simi- lar to that of Appalachian Cottontails (S. aquaticus, Kjolhaug and Woolf 1988; Citellus variegatus, Ortega 1990; Marmota flaviventris, Salsbury and Armitage 1994; Marmota monax, Swihart 1992; Sylvilagus floridanus, Trent and Rongstad 1974). We examined selection and use of home ranges at two spatial scales (i.e., macrohabitat and microhabi- tat). We quantified habitat variables within 1-m radius plots at rabbit locations and random locations both within and outside home ranges. Random plots within home ranges were located at random distances 15-35 m apart along systematically placed transects. — Random plots outside home ranges (i.e., outside areas where rabbits were located or rabbit tracks or pellets were observed) were located similarly along transects extending a distance equal to approximately one-half the diameter of the home range. Depending on the individual, the number of random plots within and outside a home range ranged from 20-38. Continuous and ordinal variables were converted to categorical variables. Habitat variables included understory stem density (0, 1-20, 21-50, 51-100, or >100 stems of any plant >0.5 m tall and <7.5 cm dbh); distance to concealment cover (< 2, 2—5, 5—10, or >10 m from a clump of understory vegetation at least 1 m? in area that provided > 50% visual obstruction: Barbour and Litvaitis 1993); percent 2002 vegetative cover (< 20, 20—< 40, 40—< 60, 60-< 80, 80-100) at 0.5-m intervals up to 1.5 m above sub- strate; percent canopy closure (0-30, 31-60, 61-80, 81-94, 95-100); percent ground cover (<20, 20—< 40, 40—< 60, 60—< 80, 80—100) of rocks, logs, and stumps; slope (< 8, 8-16, 17-24, and > 24°); and dominant (i.e., that which constituted the greatest percent cover) overstory and understory species. Percent vegetative cover up to 1.5 m was estimated using a vegetation profile board (Nudds 1977) con- structed from fabric. Percent canopy closure was estimated using a densiometer (Forest Densiometers, Arlington, Virginia). Ground cover was estimated using an ocular tube (diameter = 7.5 cm: James and Shugart 1970). Estimation of home range We measured home ranges using the minimum convex polygon (MCP: Mohr 1947) and adaptive kernel (AK: Worton 1989) methods. In addition to maximum (100%) MCP home ranges, we deter- mined areas based on utilization distributions con- taining 95% of points using MCP and AK to permit comparisons with other studies. Home ranges were represented by medians rather than means because of sample size (n = 8). We calculated home range using CALHOME (Kie et al. 1994*). CALHOME uses the Epanechnikov kernel and selects an optimum smoothing parameter that results in the lowest least-squares cross- validation (LSCV) score (Worton 1989). However, we reduced smoothing parameters to 60% of the selected value because the data were not normally distributed and this strategy prevented partitioning of home ranges into unconnected parcels (Kie et al. 1994*). We assumed locations to be independent because sufficient time (> 24 h) was allowed between loca- tions for an animal to move from one end of its home range to another (White and Garrott 1990). The num- ber of radio-telemetry locations was assumed to be adequate for determining home-range size when an STEVENS AND BARRY: HOME RANGE OF APPALACHIAN COTTONTAIL 531 asymptote was reached in a plot of home-range size against cumulative number of locations (Bowyer et al. 1995). We used a Mann-Whitney U-test (SYS- TAT® 9, Statistics I, SPSS, Inc., Chicago, IL; Zar 1999) to determine whether 100% MCP home ranges differed between males and females. Site selection and use of home range We used heterogeneity G-tests to determine whether rabbits belonged to the same statistical pop- ulation with respect to habitat. If all rabbits belonged to the same statistical population relative to any single habitat variable, we applied chi-square goodness-of-fit tests to data pooled from all rabbits (Zar 1999). These tests indicated whether habitat within home ranges differed from that outside home ranges (site selection) and whether habitat available within home ranges dif- fered from that used (1.e., at rabbit locations). When rabbits did not belong to the same statis- tical population for a habitat variable, we used log- likelihood goodness-of-fit tests (Zar 1999) to assess site selection and use of habitat. Bonferroni confi- dence intervals were then used to determine which categories of each habitat variable were selected or avoided (represented more or less often than expect- ed by chance) by individuals (Neu et al. 1974). When the expected value for a certain category of a habitat variable was <1.0, we combined values of adjacent (in magnitude) categories of the variable to prevent bias of the G-statistic (Zar 1999). All statisti- cal tests were considered significant at P <0.05. Results Home range Eight Appalachian cottontails (four males and four females) were located from 20-38 times each between February and July 1995 (Table 1). Home ranges were large and variable. Median 100% MCP home range was 5.4 ha; home-range size did not differ between the sexes (U = 2.000, p = 0.081). Median 95% MCP and AK home ranges were 4.3 and 4.7 ha respectively. TABLE 1. Home ranges (ha) of adult female (F) and male (M) Appalachian Cottontails in the Savage River State Forest, Garrett County, Maryland, as determined by 100% and 95% minimum convex polygon (MCP) and 95% adaptive kernel (AK) estimates. Number of 100% 95% 95% Rabbit locations Dates of study MCP MCP AK Fl 32 1 April-10 July 1995 2.8 2.4 3.3 F2 34 24 February-10 July 1995 1.7 1.4 1.4 F3 38 17 February-10 July 1995 6.5 5.4 5.4 F4 22 28 April-10 July 1995 4.3 3.1 2.6 MI 20 12 March-9 June 1995 6.9 5.6 5.4 M2 48 17 February 1994-10 July 1995 4.2 3.2 4.0 M3 30 3 March-10 July 1995 9.0 8.3 8.5 M4 38 17 February-10 July 1995 9.0 6.5 5.3 i —————OO——_—_—____ 582 Site selection and use of home range Considerable individual variability characterized site selection and use of home ranges. Despite this variation, we attempted to recognize common habitat associations of rabbits by noting modal responses; i.e., selection or avoidance of a certain category of a habitat variable by four or more of the eight rabbits. Rabbits were located across the entire range of elevations (730-820 m) of the study site. Eastern Cottontails occupied nearby fields, but none occurred at our study site. Heterogeneity G-tests indicated that rabbits did not belong to the same statistical population with respect to site selection, except for ericaceous and coniferous vegetation. Random locations within home ranges had no more ericaceous vegetation (a = 2539" =, 0.15. 0.50) oncaniiers (= 253, x? = 1.94, P >0.10) than random locations outside home ranges. In their selection of home ranges, rabbits avoided areas with steep slopes (>25°; Figure 1A).and those with low amounts of understory cover (<40% from 0-1.0 m and <80% from 1.1—1.5 m; Figure 1B-D). Home ranges had greater amounts of understory cover (>80% from 0—0.5 m and 1.1—1.5 m, and >60% from 0.6—1.0 m above the substrate; Figure 1B-D) than the surrounding landscape. Five rabbits (four males, one female) had some area of their home ranges bounded by agricultural fields. Some portions of the home ranges of three rabbits (one female, two males) were bounded by a stream in open, mature woodland. The home ranges of two individuals (one female, one male) were bounded, in part, by pine plantations with open understory. A steep slope (>25°) constituted some of the boundary of the home range of one male. Rabbits disproportionately used sites in close prox- imity (<2 m; Figure 2A) to concealment cover, and those with incomplete canopy closure (< 95%; Figure 2B) and R. alleganiensis as the dominant shrub species. Rabbits avoided microhabitats >10 m from concealment cover (Figure 2A), those with >95% canopy closure (Figure 2B), and those with low stem density (< 20 stems/1-m radius plot; Figure 2C). Discussion Home ranges were highly variable in size but con- sistently much greater than those previously reported for S. transitionalis (Dalke 1937). Although male home ranges were not statistically larger than those of females, power analysis (Conover and Iman 1981) indicated a low probability (@ = 0.342, P<0.30) of detecting any statistically significant differences between the sexes in home-range size because of sample size. The duration of a study could affect home-range estimates if rabbits shift their home ranges because of food availability, the distribution of potential mates, THE CANADIAN FIELD-NATURALIST Vol. 116 - or the presence of predators (e.g., White and Garrott 1990). However, we monitored rabbits for 2.5—5 months, consistent with durations of other home- range studies of Sylvilagus spp. (Chapman and Trethewey 1972; Trent and Rongstad 1974; Dixon and Chapman 1980; Kjolhaug and Woolf 1988). We did monitor rabbits during the breeding season when males travel in search of mates. However, it is unlike- ly that we overestimated home ranges because we monitored adults only, and S. obscurus may establish permanent home ranges shortly after maturity, as Forys and Humphrey (1996) found for S. palustris hefneri. We observed some home-range boundaries along edges of preferred habitat where comparatively little understory cover existed. Similarly, J. A. Litvaitis (personal communication) observed home-range boundaries of S. transitionalis consistently coincid- ing with boundaries of habitat patches in New Hampshire. Some adjacent habitats (e.g., agricultural fields) in our study would seem to constitute hard edges (Stamps et al. 1987) that inhibit movement and dispersal, illustrating how fragmentation of habi- tat could limit the distribution of S. obscurus. Rabbits also may have avoided steep slopes because traveling and resting may be difficult in such terrain. Our hypothesis that rabbits would locate home ranges in areas with the most extensive coniferous and ericaceous cover was not supported by the data. Ericaceous vegetation and conifers may be possible, but not necessary components of Appalachian Cottontail habitat. Blymer (1976) reported no erica- ceous or coniferous vegetation in a six- to seven- year-old clearcut of early successional hardwoods densely populated with Appalachian Cottontails. The habitat of New England Cottontails in New Hampshire had dense understory vegetation domi- nated by A. rubrum, Corylus cornuta, Viburnum spp., Rubus spp., and T. canadensis, but no erica- ceous species (Barbour and Litvaitis 1993; Brown and Litvaitis 1996). The presence of dense understo- ry concealment cover seems to be a more important indicator of where Appalachian Cottontails occur in western Maryland than any species of vegetation. Most ecological studies of S. transitionalis and S. obscurus have described large-scale characteristics of forests or patches of habitat (e.g., Fay and Chandler 1955; Blymyer 1976; Brewer 1980; Brown and Litvaitis 1995; Spencer and Chapman 1986). Barbour and Litvaitis (1993) found that S. transition- alis on habitat patches >5.0 ha used sites with dense understory cover and sites close to cover. Our results that Appalachian Cottontails selected home ranges with dense understory cover and used sites near con- cealment cover are consistent with these findings. Smith (1997) proposed that greater visual acuity (i.e., larger eye size and the ability to detect preda- tors at greater distances) in S. floridanus enables the Number of rabbits (of 8) i—) eo Slope (degrees) Number of rabbits (of 8) o a & <20 20-<40 % Cover at 0.6-1.0 m 40-<60 60-<80 80-100 STEVENS AND BARRY: HOME RANGE OF APPALACHIAN COTTONTAIL 533 6 a | co So 20 =)-2 5 2 -4 E z 4 8 <20 20-<40 -40-<60 60-<80 80-100 % Cover at 0-0.5m oS -_ Ss 2 = 2 = —_ i] - *) 2 E = Zz 4 -6 <20 20-<40 % Cover at 1.1-1.5m 40-<60 60-<80 80-100 FIGURE 1. Numbers of western Maryland Appalachian Cottontails (of eight total) that selected (shaded) and avoided (clear) certain categories of habitat variables (A-D) in the site selection of home ranges. Selection and avoidance were considered significant at P< 0.05. Individual rabbits may have selected more than one category. Modal responses (*) are recognized if four or more individuals selected or avoided a particular category. species to exploit open habitats where S. transition- alis (and presumably S. obscurus) suffers higher risks. Our hypothesis that S. obscurus uses home-range sites with the greatest amount of understory cover was partially supported by the avoidance of sites with low stem density. Lack of selection for high percent understory cover may have been due to two factors: (1) fairly homogeneous expanses of dense understory that provided adequate cover throughout much of the home range, and (2) differential season- al use of the home range not detected by our method- ology. These results illustrate the importance of examining habitat preferences on a variety of spatial and temporal scales. The avoidance of home-range microhabitat with low stem densities may be related to the selection of sites with R. alleganiensis. R. alleganiensis and K. latifolia were dominant in areas with high stem densities. Places with densely populated stems may be more difficult for predators to navigate. More- over, S. obscurus eats R. alleganiensis (Spencer and Chapman 1986). The evergreen K. latifolia provides cover during winter. Studies during seasons when leaves are present and absent are needed to deter- mine if seasonal variability exists in the selection of habitat by S. obscurus. Grasses and sedges make up the majority of the summer diet of S. obscurus, but woody material is the main component of its winter diet (Spencer and Chapman 1986). Sites with high percentages of canopy closure were probably avoided because understory productivity that provides necessary cover and food (e.g., Mountain Laurel, Rhodo- dendron, blueberries, and grasses) is reduced in such conditions. The considerable individual variation we found in home-range size, site selection, and microhabitat use by rabbits illustrates the importance of analyzing such variation in addition to measures of central ten- dency and interpopulation variation in traits (Hayes and Jenkins 1997; Holmes and Sherry 1997). Individuals are potentially constrained by genetics, developmental history, age, experience, and their immediate surroundings. Much of the variability in our study probably reflects variability in the immedi- ate environment (i.e., habitat) because only adults in the same area were studied. Acknowledgments This study was funded by the Heritage and Biodiversity Conservation Programs of the Maryland Department of Natural Resources, and the Frostburg State University Department of Biology. We thank the many field assistants who made this study possible, especially D. Bender and N. M. Bensley. R. P. Morgan at the Appalachian Laboratory of the University of Maryland’s Center for Environmental Studies provid- ed materials for electrophoresis, and J. Fregonara assisted with the procedure. F. L. Precht and T. C. C, Number of rabbits (of 8) <2 2-5 >5-10 >10 Distance (m) to Concealment Number of rabbits (of 8) * >100 0 1-20 21-50 51-100 Stem Density/1-m radius plot Joy provided valuable assistance with geographic equipment and software. We appreciate the support of the Savage River State Forest personnel. We thank E. Thompson and D. Feller for their valuable insights and G. Brewer, J. E. Gates, B. E. Laseter, J. R. Purdue, M. P. Scott, and an anonymous reviewer for providing helpful comments on earlier drafts. Documents Cited IUCN. 2002. 2002 IUCN red list of threatened species. www.redlist.org Downloaded 12 November 2002. Kie, J. G., J. A. Baldwin, and C. J. Evans. 1994. CALHOME home range analysis program electronic user's manual. 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Smithsonian Institution Press, Washington, District of Columbia, USA. Worton, B. J. 1989. Kernel methods for estimating the utilization distribution in home-range studies. Ecology 70: 164-168. Zar, J. H. 1999. Biostatistical analysis, 4th edition. Prentice Hall, Upper Saddle River, New Jersey, USA 663 + 212 + 11 + 20 + 23 pp. Received 23 October 2000 Accepted 16 November 2002 Distribution of Blacknose Dace, Rhinichthys atratulus, in Nova Scotia JOHN GILHEN! and ANDREW HEBDA2 Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Canada.; e-mail: !gilhenja@ gov.ns.ca; hebdaaj @ gov.ns.ca Gilhen, John, and Andrew Hebda. 2002. Distribution of Blacknose Dace, Rhinichthys atratulus, in Nova Scotia. Canadian Field-Naturalist 116(4): 536-546. The Blacknose Dace, Rhinichthys atratulus, is the most commonly misidentified minnow (Cyprinidae) in Nova Scotia. It is the only minnow native to this province having non-protractile premaxillaries. The Blacknose Dace, in Nova Scotia, is restricted to portions of The Northumberland Strait and Bay of Fundy watersheds with headwater streams originating on the Cobequid Mountains. Fifty-five localities are documented for which collections have been made. Thirty-seven addi- tional localities where no Blacknose Dace were found are also mapped. Key Words: Blacknose Dace, Rhinichthys atratulus, external characters, distribution, habitat, Cobequid Mountains, Northumberland Strait Drainage, Bay of Fundy Drainage, Nova Scotia. The Blacknose Dace, Rhinichthys atratulus (Robins et al.) [snout fish clothed in black] (Scott and Crossman 1973) is a small minnow (Cyprinidae) having an elongate, streamlined, body (Figure 1). The snout slightly overhangs the mouth (subterminal mouth). The pectoral fins are held in a horizontal position for balance in moderate flow and riffles. It is an agile swimmer adapted for living in shallow streams. The most diagnostic external character is the presence of non-protractile premaxillaries, no groove between the upper lip and snout (Figure 2). All other minnows, native to Nova Scotia, have pro- tractile premaxillaries, with a groove extending across the midline of the snout. The absence or pres- ence of a groove is easily recognizable using a hand lens. Live Blacknose Dace in Nova Scotia have a prominent, satiny-black, lateral stripe which extends around the snout, through the eyes, becomes broader on the sides of the head, continues along the trunk and caudal peduncle, ending in a blotch which streaks the base of the mid-caudal fin rays. The back is typically a light copper-orange, finely speckled with grey. The backs of a few individuals in Debert and Folly rivers, Bay of Fundy Drainage, were noted to be light iron-pyrite yellow, finely speckled with grey. This speckling extends down almost to the black stripe, thus leaving a narrow, lustrous, copper- orange, or iron pyrite-yellow, stripe bordering the black stripe. One very dark adult male from Pugwash River was black on the back, which was continuous to the black stripe and lacking a lustrous border. The lower sides are pearly-white becoming silvery-white on the belly. Sexually mature males display a reddish blush on the fins Gilhen 1974, particularly the pectoral fins. When males are in breeding condition and courting females, the red colour becomes so intense that the black lateral stripe is no longer discernable. However, within a few minutes of a male being removed from the spawning site and placed in an aquarium it regains the black stripe leaving only the fins and sides blushed in red. The average total length of adult Blacknose Dace collected in Nova Scotia is 58.5 millimeters, ranging from 50 to 72 mm (n=71). Scott and Crossman (1973) gave the average length as 64 mm. Distribution The Blacknose Dace is found in eastern and cen- tral Canada from northwestern mainland Nova Scotia, west through New Brunswick, southern Quebec and Ontario, in Lake of the Woods but not northeast of Lake Superior, northwest to central Manitoba; and southward in the United States, on both sides of the Appalachian Mountains, to Georgia, Alabama and Mississippi (Scott and Cross- man 1973; Coad et al. 1995). In Nova Scotia, R. atratulus is restricted to water- sheds with headwater streams on the Cobequid Mountains of the northern mainland (Figure 3). The streams on the north side, which empty into the Northumberland Strait, where we found Blacknose Dace are (from west to east) Shinimicas River (SHR), River Philip (RP), Pugwash River (PUR), Wallace River (WR), French River (FR), Waughs River (WAR) and River John (RJ) (Appendix 1). Streams on the south side which empty into the Bay of Fundy are, from west to east, Maccan River (MAR), Hebert River (RH), Moose River (MR), Portapique River (PR), Great Village River (GVR), Folly River (FR), Debert River (DR), North River (NR) and Salmon River (SR) (Appendix 2). Habitat The Blacknose Dace in Nova Scotia inhabits only those watersheds having first order streams draining the north or south faces of the Cobequid 536 2002 GILHEN AND HEBDA: BLACKNOSE DACE IN NoVA SCOTIA 357 FiGurE 1. A computer-enhanced photograph of a living adult male Blacknose Dace, Rhinichthys atratulus, from Christie Brook, a tributary of Salmon River, Bay of Fundy Drainage, Nova Scotia, 7 May 1998 (Negative Number-24,377 frame # 20). Mountains. Natal rivers and some of their tribu- taries, typically have extensive shallow sections with riffles. The clear, cool water flows at a moder- ate rate over gently-sloping hard bottom. R. atratu- lus shares this specific habitat with Brook Trout, Salvelinus fontinalis, in brooks (Figure 4) and Atlantic Salmon parr, Salmo salar, in rivers (Figure 5). The described habitat helps to explain why this no groove between upper lip and snout subterminal mouth Blacknose Dace ' non-protractile premaxillary complete groove between upper lip and snout \ FW. Scott 2000 protractile premaxillary FiGuRE 2. Sketch showing non-protractile premaxillaries of a Blacknose Dace, Rhinichthys atratulus, from Christie Brook, a tributary of Salmon River, Bay of Fundy Drainage, Nova Scotia (NSM12635 (.1)) and protractile premaxillaries of a Lake Chub, Couesius plumbeus, from Paradise Brook, a tributary of Tidnish River, Northumberland Strait Drainage, Nova Scotia (NSM12665 (.1)). stream fish is not found in all watersheds originat- ing on the Cobequid Mountains. We did not find Blacknose Dace in streams which cascade or in slow, deep, soft-bottomed sections, such as exten- sive still-waters or in parts which meander through bogs. When we checked marginal waters, such as quiet shallow inlets and at the mouths of slow tribu- taries, of streams where R. atratulus were found we observed mixed schools of cyprinids including juveniles of Blacknose Dace, Common Shiner (Luxilus cornutus), Northern Redbelly Dace (Phoxinus eos), and Creek Chub (Semotilus atro- maculatus), and other freshwater species such as juvenile White Sucker (Catostomus commersoni), and Threespine Stickleback (Gasterosteus aculea- tus). When we checked streams at the periphery of the range of Blacknose Dace, such as Paradise Brook (Tidnish River watershed), and West River (Pictou watershed), where no Blacknose Dace were found, the niche was occupied by Lake Chub, Couesius plumbeus. The mis-named Lake Chub is also an elongate streamlined minnow well-adapted to living in shallow streams. This could explain why juvenile Lake Chub have been mis-identified as Blacknose Dace. Discussion In distribution, the Blacknose Dace is one of the most restricted of the freshwater fishes native to Nova Scotia. Although it can be identified by check- ing the non-protractile premaxillaries, it is the most commonly misidentified minnow in this Province. This is a result of the lack of attention to non-game fishes during fish surveys for environmental assess- ments. In Nova Scotia, when such faunal surveys are undertaken, there is no requirement for the collection of voucher specimens of non-game fishes or their deposition in a repository collection. The Nova Scotia Museum of Natural History has an extensive fish collection and is one institution where environ- 538 We vminvindsae © fe enccnss Bees cede Ao SR. A. Af... BU THE CANADIAN FIELD-NATURALIST PR GV TT ———— Ec imo NM Vol. 116 @ present not present fn gh cee esis oem Ot ees ETE: Me ree terre ed aes eens thea cae ey sees seus sewsny Meser eye err i q Lak R FR DR NR SR FiGurE 3. Distribution of Blacknose Dace, Rhinichthys atratulus, in Nova Scotia. Not Present, represented by open circles on the map, means localities where Blacknose Dace were looked for but not found. In the Northumberland Strait Drainage the letters SHR refers to Shinimicas River; RP, River Philip; PUR, Pugwash River; WR, Wallace River; FR, French River; WAR, Waughs River; and RJ, River John (see Table 1). In the Bay of Fundy Drainage, MAR refers to Maccan River; RH, River Hebert; MR, Moose River; PR, Portapique River; GVR, Great Village River; FR, Folly River; DR, Debert River; NR, North River and SR, Salmon River, (see Table 2). mental assessment reports are reviewed for com- pleteness and accuracy and the identification of specimens can be verified. Errors in identification of Blacknose Dace date back to 20 July1941 when C. F. Allen mis-identified three Northern Redbelly Dace, P. eos, from Grand Lake, Halifax County, as Striped Dace, Rhinichthys atronasus, the common and scientific names in use at that time for Blacknose Dace (Scott and Crossman 1973). This record was repeated by Davis (1981). There are a number of other incorrect records, also from localities outside the range for Blacknose Dace in Nova Scotia. We suspect Blacknose Shiner, Notropis heterolepis, have been incorrectly identi- fied as Blacknose Dace in a report on Lake Egmont (Parker 1994). Another such record is Anonymous (1998) in which Blacknose Dace was reported from 35 localities across the northern mainland; 11 are in north-eastern Nova Scotia well outside its con- firmed range. Between 1972 and 2000, J. G. investi- gated streams between West River Pictou, Pictou: County; and St. Marys River, Guysborough County, for freshwater fishes and did not find Blacknose Dace at any of the sampled sites. Juvenile Creek Chub were observed in the St. Marys River that have a prominent black lateral band which extends onto the head, possibly accounting for previous con- fusion. During a study of the food of mergansers, White (1957) did not find Blacknose Dace in the stomachs of these fish-eating birds from three streams in the northeast mainland including Moser River, East River St. Marys and West River St. Marys. He did, however, find Blacknose Dace in stomachs of mer- gansers at Wallace River (WR), Maccan River (MAR), River Hebert (RH) and River Philip (RP) of the Carboniferous area of mainland Nova Scotia where we also found R. atratulus (Figure 3). Blacknose Dace were first collected in Nova Scotia at Higgens Brook, a tributary of Wallace River (WR), by A. G. Huntsman on 30 July 1937 (Appendix 1). The second collection of R. atratulus for this Province was at McPherson Bridge, Shinimicas River (SHR), by an unknown collector, 13 September 1944. Both collections are from Northumberland Strait drainage of Nova Scotia and are in the Royal Ontario Museum (ROM), Toronto. Dan A. Livingstone collected freshwater fishes in FIGURE 4. Christie Brook, a tributary of Salmon River, Bay of Fundy Drainage, Nova Scotia, habitat where Blacknose Dace, Rhinichthys atratulus, White Sucker, Catostomus commersoni, Brook Trout, Salvelinus fontinalis, and Threespine Stickleback, Gasterosteus aculeatus, were recorded. Photograph taken on 13 August 1998 (Negative Number 24,379: Frame Number 2). Nova Scotia over the four-year period 1947 to 1950. He also visited the Royal Ontario Museum and noted the collections of freshwater fishes from Nova Scotia. He did not find the two earlier samples of R. atratulus. These records are not mentioned by Scott and Crossman (1973) and may have been deposited ROM at a later date. Livingstone (1952), therefore, lists only the two localities, Shinimicas River (SHR) FiGureE 5. Salmon River at road to Riversdale, Bay of Fundy Drainage, Nova Scotia, habitat where Blacknose Dace, Rhinichthys atratulus, White Sucker, Catostomus commersoni, young Rainbow Smelt, Osmerus mordax, Atlantic Salmon parr, Salmo salar and Threespine Stickleback, Gasterosteus aculeatus, were recorded. Photograph taken on 13 August 1998 (Negative Number 24,380: Frame Number 8). GILHEN AND HEBDA: BLACKNOSE DACE IN NOVA SCOTIA 539 and River Philip (RP), for Blacknose Dace which he discovered during his 1950 survey. In the summer of 1955 Francis R. Cook and John S. Erskine collected fishes in lakes and streams as they criss-crossed the mainland of Nova Scotia and added five new localities for Blacknose Dace. A collection of two specimens from Atkinson Brook in the River Hebert (RH) watershed on 30 July 1955 is the first record of R. atratulus for the Bay of Fundy Drainage (Cook 1961) (Appendix 2). A collection of one speci- men on | August 1955 from a stream near Dewar (Angevine) Lake is believed to be from Flemming Brook, a small tributary of Wallace River (WR). Most samples of Blacknose Dace in the fish col- lection of the Nova Scotia Museum of Natural History were collected between 1964 and the pre- sent, with the greatest sampling effort carried out by us during spring and summer of 1997 and 1998. A beach seine and pond dip net were effective in col- lecting specimens. After four years collecting and observing freshwa- ter fishes in Nova Scotia, Livingstone (1952) noted the largest number of species were found in Cumberland County and that the number of species diminishes to the east and south through the Province. He hypothesized that “This lack of range discontinuities indicates that complete destruction of the pre-Pleistocene fish fauna occurred during the glaciation of Nova Scotia, and that the species now here have repopulated the province since the reces- sion of the ice sheet. The species abundance in the various regions indicates that they entered Nova Scotia by way of the Isthmus of Chignecto, not by some [southern] land bridge no longer existing.” The distribution of the Blacknose Dace is one example used to support this hypothesis. The post-glacial re- colonization of lakes and streams in Nova Scotia, however, is probably more complex than previously thought, and may involve westward migrations from Atlantic refugia (Howden et al. 1970; Marcogliese 1992; Nedeau et al. 2000). Because recognition of distinct populations and peculiar distribution patterns of freshwater fishes in Nova Scotia probably holds clues to the post- Wisconsin Glaciation re-colonization in this Prov- ince it is important that we have a clear understand- ing of the distribution of each species in this Province. Today, however, the integrity of these interesting freshwater fish distribution patterns, par- ticularly of forage fishes, such as minnows, is in jeopardy due to the introductions of foreign, predato- ry, freshwater game species, notably Chain Pickerel, Esox niger, and Smallmouth Bass, Micropterus dolomieu. Also, as Livingstone (1952) warned: “Similarly the salmonids will probably disappear from many of the marginal waters in which they now occur when faced with the competition of pikes and centrarchids”. 540 Acknowledgments The authors are grateful to Roger Lloyd for pho- tographing Blacknose Dace habitat and live speci- mens in an aquarium. We thank Sylvie Laframboise, Canadian Museum of Nature, Erling Holm, Royal Ontario Museum and Lawrence K. Benjamin, Nova Scotia Department of Natural Resources, for providing data on collections in their respective institutions. We thank Karen March and John MacMillan, Porter Dillon Limited, Halifax; Alfonso Rojo, St. Marys University, Halifax and Barry Sabean, Nova Scotia Department of Natural Resources, Kentville for the donation of specimens. We thank John Chapman, Edward Claridge, Rebecca Fleming, Karen Gilhen, Michael Gilhen, Jennifer Russell, Fred Scott and Jason Taylor for their assistance in field work. We also thank Fred Scott, curator, Acadia University Wildlife Museum, Wolfville, for allowing us access to the fish collec- tions in that institution. Fred Scott produced Figures 2 and 3 and Karen Gilhen Appendicies 1 and 2. Leslie Pezzack read and made a number of useful comments on the manuscript. Collecting per- mits were provided by Greg Stevens, Senior Advisor, Anadromous & Freshwater Fisheries Resource Management Branch, Scotia-Fundy Fish- eries, Department of Fisheries and Oceans, Halifax, Nova Scotia. Literature Cited Anonymous. 1998. Maritimes & Northeast Pipeline. Environmental Protection Plan (Mainline), Revision 01. Halifax. Coad, B. W., H. Waszczuk, and I. Labignan. 1995. Encyclopedia of Canadian Fishes. Canadian Museum of Nature and Canadian Sportfishing Productions Inc. Ottawa. 928 pages. Cook, F. R. 1961. Further records of the eastern black- THE CANADIAN FIELD-NATURALIST Vol. 116 nose dace in Nova Scotia. Canadian Field-Naturalist. 75: 51-52. Davis, D. S. 1981. Aquatic fauna of the Shubenacadie headwater lakes. An exercise in information retrieval. Curatorial Report Number 44. Nova Scotia Museum.15 pages. Gilhen, J. 1974. The fishes of Nova Scotia's lakes and streams. Nova Scotia Museum, Halifax. 49 pages. Howden, H. F., J. E. H. Martin, E. L. Bousfield, and D. E. McAllister. 1970. Fauna of Sable Island and its zoogeographic affinities — A compendium. National Museum of Natural Sciences. Publication in Zoology (4): 45 pages. Livingstone, D. A. 1952. The freshwater fishes of Nova Scotia. Proceedings Nova Scotian Institute of Science 23: 1-90. Marcogliese, D. J. 1992. First report of the Threespine Stickleback, Gasterosteus aculeatus, from Sable Island. Canadian Field-Naturalist. 106: 264-266. Nedeau, E. J.. M. A. McCollough, and B. I. Schwartz. 2000. The freshwater mussels of Maine. Maine Department of Inland Fisheries and State House Station Number 41. Augusta, Maine. 118 pages. Parker, E. 1994. Lake Egmont: Survey of flora and fauna of an unusual eco-system. Halifax, Nova Scotia. 77 pages. Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott. 1991. Common and Scientific Names of Fishes from the United States and Canada. Fifth Edition. American Fisheries Society. Special Edition 20: 183 pages. 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Received 27 December 2000 Accepted 23 December 2002 541 BLACKNOSE DACE IN Nova SCOTIA GILHEN AND HEBDA 2002 panuljuoo ‘OD purjoquing M.OL ES 0£€9 (du) ‘uonouNs Ps1OJXO ¢ Us I) UYor 91D'easI ‘NuSE TP SP dipiyd Joary ye dipiyd J9Ary [1 1-Z-P86INSN P86l Aine g1 MST iE€ 0&9 CUM) OD) puejroquin’) 6 asp “V WoL OLW'PV7Z "NuSP 6€ oSP JOATY 9OBTTE AA “TOATY 9ORTTR AA OF l-PBOWN ER61 Aine g M.S iTS o£9 (du) OD) puejroquin’) I] aspy ‘Vv WoL, TIH £81 *"N..80 VP oSP dipiyd JOAry “JOATY O[NV] 67V I -PBONN €R61 Aine g MSP LS o£9 (du) ‘OD puefrequin’) | aspq “V WoL, [TH'€asl ‘N..80 PP oSP dipiyd Joary ‘JOATY OPV] 87V1-P8ONN ER6I Aine g M.S 1S o£9 (du) OD) pue[joquin’) | MNO Xe] *N..80 VP oS? dipryd JoAry ‘dipiyd J9Ary OOSTIWSN LL61 eune {I USYTID) JSBYI|| ‘OD puejsoquin’) ‘ueuideyD uyor MN. OP £9 (UM) “TOATY SORTER AA Ol ‘usy ID UYyor TIN PHsl *"N..OS i8E oSP JOATY 9ORTTE AA Yourlg SoA L-Z-VI6BINSN p96! 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STANIFORTH!, WILLIAM J. Copy2, and KATHERINE A. FREGO? 'Department of Biology, University of Winnipeg, 515 Portage Ave., Winnipeg, Manitoba R3B 2E9, Canada; e-mail: richard. staniforth @uwinnipeg.ca Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-food Canada, Crop Protection Program, William Saunders Bldg., Central Experimental Farm, Ottawa, Ontario K1A 0C6, Canada *Department of Biology, University of New Brunswick, P.O. Box 5050, St. John, New Brunswick, E2L 4L5, Canada: e-mail: frego@unbsj.ca Staniforth, Richard J., William J. Cody, and Katherine A. Frego. 2002. Bill Dore’s notes on the Kaladar cactus (Opuntia fragilis). Canadian Field-Naturalist 116(4): 547-550. The late botanist William G. Dore habitually left copious notes with his herbarium specimens and kept files on interesting plants while at the Biosystematics Research Institute of the Canada Department of Agriculture in Ottawa, Ontario. Analysis of his specimens, notes and letters regarding an isolated colony of Brittle Prickly-pear Cactus (Opuntia fragilis) from Kaladar in eastern Ontario led to the unfolding of a botanical story of discovery and rediscovery. This colony was first discovered in 1934, the location apparently lost and subsequently rediscovered in 1947, more or less neglected, and then brought to the attention of a wider circle of botanists in the mid 1960s. Attempts to determine its taxonomic identifi- cation were thwarted by the infrequency of flowering until the early 1960s. Key Words: William (Bill) G. Dore, Brittle Prickly-pear Cactus, Opuntia fragilis, discovery, determination, Kaladar, Ontario. The eminent Canadian plant biologist and natural- ist, William (Bill) G. Dore, left a legacy of botanical contributions when he died in 1996 (see Darbyshire 1998). These included excellent herbarium speci- mens in the Vascular Plant Herbarium of the Biosystematics Research Institute in Ottawa (now called the Eastern Cereal and Oilseed Research Centre, Agriculture and Agrifood Canada), to which supplemental information had been routinely appended (Darbyshire 1998). Bill Dore’s specimens with their added notations and his cactus files showed a particular personal interest in a remarkable disjunct colony of Opuntia fragilis (Brittle Prickly- pear Cactus) from a single rock outcrop, near Kaladar in eastern Ontario (see Beschel 1967b; Staniforth and Frego 2000). This population is approximately 1000 km east of the nearest part of the main range for the species, in Wisconsin (see dis- tribution map in Staniforth and Frego 2000). The objective of this note is to explore Bill Dore’s speci- mens, annotations, notes and letters on the Kaladar cactus in order to document the events surrounding its discovery, determination and possible origin. Materials and Methods Relevent specimens, letters, annotations and notes were obtained from two sources. Firstly, Dore gener- ously provided a copy of his entire Opuntia file to K.A.F. in 1980 (Frego 1980*, Dore 1980*). Secondly in April 2000, W.J.C. retrieved materials from the herbarium file for Opuntia fragilis housed in the Vascular Plant Herbarium of the Eastern Cereal and Oilseed Research Centre, Agriculture and Agrifood Canada. Together, these consisted of four herbarium specimens (MacClement 6372, Dore and Senn 47- 671, Moore 2765, Dore 22811) with their annotation labels, 11 letters sent by or to Bill Dore, two pages of typed and four pages of hand written notes. The speci- mens and written material spanned the years from 1934 to 1980. Collectively these items captured the history of the discovery of the colony, the determina- tion of identification for the cactus species, and some of the thought that went into the consideration of the colony’s origin. We organised photocopies of the items into a chronological sequence (see Documents and Specimens Cited section) and their information contents were linked to present a history of discovery for Opuntia fragilis in eastern Ontario. Results and Discussion Discovery Local people knew of the cactus colony long before it was “discovered” by a professional botanist in 1934. According to Mary I. Moore of Deep River, Ontario (Moore 1965*, 1967*, 1973*), three genera- tions of the Smedley family had known of its loca- tion since about 1905. On the other hand, and to the frustration of the researchers, directions to the site by locals usually turned out to be erroneus. For instance, David Brydson, the owner of a local hotel, was notably inaccurate regarding the colony's whereabouts and its flowering characteristics, even though he had transplants growing in his own garden (Dore 1947*, 1949b*,1967c*). W. T. MacClement (Professor of Botany, Queen’s University, Kingston 1906-1936; see Smallman et al. 1991) appears to have made the first botanical collec- tion of the cactus in May 1934 (MacClement 1934*). 547 548 We have no information as to how MacClement became aware of the site. Claude Garton (Lakehead University) later recalled (Dore 1978*) that the event had been a Sunday field trip to the site for botany stu- dents from Queen’s University. A pressed specimen was sent to Merritt Fernald at the Gray Herbarium, Harvard University, for identification. However, there is no record of any determination having been made by Fernald and it appears that the specimen was overlooked for the next 31 years. The MacClement field trip also resulted in living plants being cultured in the greenhouse at Queen’s by MacClement and his successors R. O. Earl (Dore 1967c*) and A. Crowder (Crowder 1973*). Apparently, unaware of the MacClement “discov- ery” and the existence of the Harvard specimen (Dore 1967a*), Dore and others continued trying to verify rumours that a wild cactus existed somewhere in southeastern Ontario (Dore 1947*). Herbert Groh (Department of Agriculture, Ottawa) and Dore, act- ing on erroneus directions by local people, searched the Kaladar area without success but may even have been as close as the opposite side of the road in 1942 (Dore 1947*). Jack Gillett eventually relocated the colony on 30 July 1947 (Dore 1947*) and informed Dore, who travelled to the site with Harold Senn dur- ing the following week and procured a specimen for THE CANADIAN FIELD-NATURALIST = : Vol. 116 - the Biosystematics Research Institute (Dore and Senn 1947*). The site was next visited by Dore, Senn and Clarence Frankton (Department of Agriculture, Ottawa) on 17th August 1949 (Dore 1949b*). The next recorded visit did not occur until 3rd May 1965 (Moore 1965*), when Mary Moore (Deep River, Ontario) collected a specimen. Coinciden- tally, in June 1965, Bernard Boivin of Agriculture Canada unexpectedly encountered the forgotten MacClement specimen while visiting the Gray Herbarium. He reported its existence to his colleague Bill Dore (Dore 1967a*). It appears that no one in Ottawa was aware of the existence of this early speci- men, and its discovery must have been quite an excit- ing event. A photograph of the original Gray Herbarium specimen was soon accessioned into the Herbarium of the Biosystematics Research Institute (MacClement 1934*). This was likely to have been the stimulus behind renewed investigations into the cactus by Dore and others in the mid- and late 1960s (Dore 1966*). On 30th May 1967, Dore, Roland Beschel (Queen’s University, Kingston), Paul Maycock (Erindale College), Gunnar Wassen (National Herbarium, Ottawa) and probably others visited the site. Wassen took black-and-white photographs of the plants (Dore 1967b*) and the site (Figure 1). sl Coes FiGuRE 1. A photograph of the Brittle Prickly-pear Cactus (Opuntia fragilis) site, near Kaladar, southeast Ontario taken by Gunnar Wassen during a field trip with W. Dore, R. Beschel, P. Maycock (and possibly others) in May 1967. 2002 FIGURE 2. A photograph of cactus cladodes “attached to Paul Maycock’s big boot” (Dore 1967b) taken by G. Wassen in May 1967 at the Kaladar site. Later that same year, Beschel led the Canadian Botanical Association on one of its first field trips which included a visit to the cactus colony (Beschel 1967a*). In June 1967, Beschel published a descrip- tion of the site and the cacti in the Quarterly Bulletin of the Kingston Naturalists (Beschel 1967). At the same time and perhaps unwittingly, Dore drafted a longer article and sent it to Beschel for review and possible co-authorship (Dore 1967c*). In his cover- ing letter to Beschel, Dore recommended that an appropriate place for publication would be the Ontario Naturalist. This article was never published even though Beschel revisited the site in July 1967 to take more photographs and gather more descrip- tive information about the cacti and other plants at the site (Beschel 1967b*). Determination The identity of the Kaladar cactus appears to have been nearly as elusive as its discovery. This was like- ly attributable to its similarity to certain other species in the genus (such as O. polyacantha) and to the rari- ty of flowers in Kaladar plants. The MacClement specimen was originally determined as “Opuntia” in 1934 (MacClement 1934*). R. O. Earl of Queen’s University sent the MacClement specimens to M. O. Malte (Chief Botanist, National Museum) for identi- fication but the lack of flowers led to inconclusive results (Dore 1947*). During 1949, Dore wrote to cactus experts Elzada Clover (University of Michigan) and H. A. Shetrone (Ohio State Museum) to seek help with the identification process (Dore 1949a*, 1949c*; Meyer 1949*) but there is no docu- STANIFORTH, CoDy, AND FREGO: KALADAR CACTUS 549 mentation of any reply. In the 1950s and early 1960s, most specimens (Dore and Senn 1947*; Moore 1965*; Dore 1966*) were being labelled as “Opuntia (?fragilis) Haw.” but the origin of this tentative deter- mination is unknown as is the basis on which it was made. There seems to have been no uncertainty in the determination by the time that Beschel undertook his studies on the cactus in 1967 (Beschel 1967a*, 1967b*). All specimens that we examined had been confirmed with annotations as Opuntia fragilis (Nutt.) Haw. by Lyman Benson (Pomona College, Claremont, California) in April 1972 and by Bruce Parfitt (Arizona State University) in 1991. Colony origins Alternate hypotheses to explain the origin of this anomalous disjunct distribution have been developed by Beschel (1967b*), Dore (1967c*) and Staniforth and Frego (2000). The two hypotheses are: (1) human introduction at an unknown date, or (2) as a relict of a wider distribution during hypsithermal period, 8000-4000 years before present. Dore’s doc- uments in the late 1940s indicate that these hypothe- ses were already being formulated at that time; i.e., “Mr. Gillett is of the opinion that the cactus is introduced or escaped” (Dore 1947). At the Annual General Meeting of the Canadian Botanical Association (Vancouver, June 1980), Dore hypothe- sised that both Lithospermum canescens and Opuntia fragilis may have been purposely spread by aboriginal people because the former possesses a red dye whereas mucillage from stems of the cactus can act as a mordant for the dye. We were recently enlightened by Daniel Brunton of Ottawa (Brunton 2000*) of another of Dore’s explanations: “The col- onization road upon which the nearby highway was later constructed was put in when a 19th century gold rush occurred in the area just to the north (of Kaladar); this was, I believe, about the time that the California gold rush was winding down. It’s not hard to imagine some western gold seeker unknowingly brought a pad along on his gear, only to have it fall or be broken off at the Kaladar site”. In support of the relict hypothesis, Dore wrote that he, Clarence Frankton and Harold Senn had observed other asso- ciated species at Kaladar “which may indicate affini- ty with a more southern or western climatic area” (Dore 1949b*) suggesting that the cactus was part of a relict hypsithermal community. This second hypothesis is now thought to be less likely (Staniforth and Frego 2000), largely because closely associated species do not have western or southern affiliations as was originally proposed. However, some southern species reach their northern limits in the general region (Brownell et al. 1996). Conclusions The isolation of disjunct plant populations intrigued and challenged Bill Dore’s imagination for explanations. The Brittle Prickly-pear Cactus 550 growing near Kaladar, in eastern Ontario: a popula- tion located more than 1000 km from the main range of the species is an excellent example of this. We can imagine that rumours of its existence amongst local people reached Bill Dore and other Ottawa (and Kingston) botanists, in the 1940s. The task of finding the colony (and perhaps the challenge of who would be the first to detect it!) resulted in several botanising expeditions, but it was Jack Gillett who eventually discovered the site. The hunt was then replaced by the challenge of formulating an accept- able, logical, ecological or historical explanation of its origins. Dore excelled at this. Darbyshire (1998) noted that “Bill is well known for his hypotheses of aboriginal and early European influences” when attempting to explain the unusual distributions of plants, but cautions, “Although many people have been quick to reject these explanations of present- day plant distributions, few have offered contesting hypotheses to explain ‘how’ and ‘why’”. Dore’s let- ters, specimens and copious annotations allowed us to follow the events which led to the discovery, species determination and hypotheses of origin for this unusual population and in so doing, they have contributed an interesting aspect to the history of dis- covery of the Canadian flora (see Pringle 1995). Acknowledgments The authors are grateful to Mrs. R. E. Beschel of Kingston for providing us with the photographs that had been taken during the 1967 field trip to the cac- tus site near Kaladar, Ontario. These photographs and also the documents mentioned below are filed with R.J.S. at the University of Winnipeg. Documents and Specimens Cited (marked * in text), in chronological sequence MacClement, W. T. 1934. Photograph of Herbarium specimen of Opuntia fragilis. Accessioned from Gray Herbarium. Collection number: MacClement 6372 (DAO 269655). Dore, W. G., and H. A. Senn. 1947. Herbarium specimen of Opuntia fragilis. Collection number: Dore & Senn 47-671 (DAO 82191). Dore, W. G. 1947. Two typed pages, signed and dated (“Fall 1947”) and identified as “Re. 47.671”. Attached to Collection number: Dore & Senn 47-671 (DAO 82191). Dore, W. G. 1949a. Letter to Dr. Bernard S. Meyer, Ohio State Unversity. Dated 4 April 1949. Meyer, B.S. 1949. Letter to W. G. Dore. Dated 14 April 1949. Dore, W. G. 1949b. Two hand-written pages, signed and dated (“Dec. 5, 1949”). Attached to Collection number: Dore & Senn 47-671 (DAO 82191). THE CANADIAN FIELD-NATURALIST 2, | Vol. 116 - Dore, W. G. 1949c. Letter to Dr. Elzada Clover, University of Michigan. Dated 6 December 1949. Moore, M. I. 1965. Herbarium specimen of Opuntia frag- ilis. Collection number: Moore 2765. (DAO 82194) Dore, W. G. 1966. Herbarium specimen of Opuntia frag- ilis. Collection number: Dore 22811. (DAO 82193) Dore, W. G. 1967a. Handwritten note dated (18 May 1967) and signed. Attached to MacClement, W. T. 6372 (DAO 269655). Beschel, R. E. 1967a. Field trip 1 to the Kingston region led by R. E. Beschel, Queen’s University. Annual meet- ing of the Canadian Botanical Association. Ottawa. 30 May 1967. Mimeographed sheets. Dore, W. G. 1967b. Letter to Dr. Gunnar Wassen. Dated 15 June 1967. Dore, W. G. 1967c. Letter to Roland E. Beschel. Dated 18 June 1967. Attached to a rough draft of a 12 page manuscript “Rare Cactus in Ontario Wilderness by William G. Dore and Roland E. Beschel”. Beschel, R. E. 1967b. Letter to W. G. Dore. Dated 27 July 1967. Moore, M. I. 1967. Letter to W. G. Dore. Dated 17 August 1967. Moore, M.I. 1973. Letter to W. G. Dore. Dated 9 April 1973. Crowder, A. 1973. Letter to W. G. Dore. Dated 17 April 1973. Dore, W. G. undated (but post-1978). Hand-written note regarding visit to cactus area by Claud Garton and Prof. MacClement in 1933 or 1934. Dore, W. G. 1980. Letter to Katherine Frego. Dated 2 July 1980. Frego, K. A. 1980. A copy of Dore’s “cactus file” given to K. A. Frego in 1980. Brunton, D. 2000. Letter to R. J. Staniforth. Dated 20 July 2000. Literature Cited Beschel, R. E. 1967. The cactus at Kaladar. The Bluebill: Quarterly Bulletin of the Kingston Field Naturalists 14: 11-12 Darbyshire, S. J. 1998. A tribute to William George Dore, 1912-1996. Canadian Field-Naturalist 112: 357-365. Pringle, J.S. 1995. The history of the exploration of the vascular flora of Canada. Canadian Field-Naturalist 109: 291-356. Smallman, B. N., H. M. Good, and A. S. West. 1991. Queen’s Biology: an academic history of innocence lost and fame gained, 1858-1965. Queen’s University Press. Kingston, Ontario, Canada. 215 pages. Staniforth, R. J., and K. A. Frego. 2000. Ecological his- tory and population dynamics of a disjunct population of Brittle Prickly-pear Cactus, Opuntia fragilis (Cactaceae), in eastern Ontario. Canadian Field-Naturalist 114: 98-105. Received 20 December 2000 Accepted 14 February 2003 Amphibian and Reptile Diversity along the St. Lawrence River BENO!T JoBIN!, DAVID RODRIGUE?, and JEAN-LUC DESGRANGEsS! ‘Canadian Wildlife Service, Environment Canada, 1141 Route de l’Eglise, C.P. 10100, Sainte-Foy, Québec G1V 4H5 Canada St. Lawrence Valley Natural History Society, 21125 chemin Sainte-Marie, Sainte-Anne-de-Bellevue, Québec H9X 3Y7 Canada Jobin, Benoit, David Rodrigue, and Jean-Luc DesGranges. 2002. Amphibian and reptile diversity along the St. Lawrence River. Canadian Field-Naturalist 116(4): 551-558. The Biodiversity Portrait of the St. Lawrence project was initiated to gain insight into the natural and anthropogenic factors that affect biodiversity and to have a better understanding of species distribution along the St. Lawrence River. This paper presents a regional analysis of the diversity of herpetofauna living along the Québec portion of the St. Lawrence River and identifies areas to be protected for the maintenance of amphibian and reptile populations in southern Québec. Data collect- ed since the end of the 19" century were compiled in geographic units (natural provinces) and in a systematic grid of 100- km? quadrats. The herpetofauna of the St. Lawrence River is composed of 33 species: 18 amphibians and 15 reptiles. The most commonly recorded amphibians were the American Toad (Bufo americanus), Spring Peeper (Pseudacris crucifer), Northern Leopard Frog (Rana pipiens) and Wood Frog (Rana sylvatica), and the most abundant reptiles were the Common Garter Snake (Thamnophis sirtalis), Painted Turtle (Chrysemys picta) and Common Snapping Turtle (Chelydra serpentina). Species richness was greater in the southern portion of the St. Lawrence in the Montréal region where most records of species with high-priority conservation status in Québec were also concentrated, making this a high-priority region for conserving the biodiversity of Québec’s herpetofauna. Key Words: amphibians, reptiles, herpetofauna, St. Lawrence, biodiversity, conservation, species richness, distribution. La production du Portrait de la biodiversité du Saint-Laurent visait 4 mieux comprendre les facteurs naturels et anthropiques qui agissent sur la biodiversité et a préciser nos connaissances sur la distribution des espéces fauniques et floristiques vivant le long du fleuve Saint-Laurent. Nous présentons une analyse régionale de la diversité de | herpétofaune retrouvée dans la portion québécoise du Saint-Laurent et nous identifions des secteurs ol des actions de conservation devraient étre entreprises afin de protéger les populations d’amphibiens et de reptiles du sud du Québec. Les observations compilées depuis la fin du 19°™* siécle ont été intégrées dans un cadre écologique (provinces naturelles) et dans une grille arbitraire formée de parcelles de 100 km’. L’herpétofaune du Saint-Laurent est composée de 33 espéces soit 18 amphibiens et 15 reptiles. Les amphibiens les plus fréquemment observés sont le Crapaud d’ Amérique (Bufo americanus), la Rainette crucifére (Pseudacris crucifer), la Grenouille léopard (Rana pipiens) et la Grenouille des bois (Rana sylvatica), tandis que les reptiles les plus communs sont la Couleuvre rayée (Thamnophis sirtalis), la Tortue peinte (Chrysemys picta) et la Chélydre serpentine (Chelydra serpentina). La richesse spécifique était plus élevée dans la région de Montréal, 1a ot la majorité des observations des espéces ayant un statut de conservation élevé ont également été effectuées. Cette région présente donc un intérét majeur pour la conservation de la biodiversité de l’herpétofaune québécoise. Mots-clés : amphibiens, reptiles, herpétofaune, Saint-Laurent, biodiversité, conservation, richesse spécifique, distribution. The St. Lawrence Vision 2000 Action Plan (SLV bution of the major taxonomic groups, from plants to 2000), a federal-provincial program developed to mammals, living in or along the St. Lawrence River study and protect the St. Lawrence River ecosys- (DesGranges and Ducruc 2000*). tems, was implemented to gain insight into natural Amphibian and reptile distribution was part of this and anthropogenic factors that affect biodiversity portrait, being of particular interest considering the and to have a better understanding of species distri- declining population trends of several species, espe- bution along the St. Lawrence River, in order to cially in anurans (frogs and toads), recently observed direct conservation efforts in areas where species at in Canada (Bishop and Pettit 1992; Green 1997) and risk are concentrated. Because the different sources worldwide (Blaustein and Wake 1990; Phillips of information on the physical and biotic characteris- | 1990). Because many amphibian and reptile species tics of the St. Lawrence were scattered in the scien- rely on wetlands or riparian habitats to complete tific literature, in unpublished reports, and in non- their life cycle, they are prone to be greatly affected centralized databases, the Biodiversity Portrait of the by wetland drainage and habitat losses. It is estimat- St. Lawrence project was initiated in 1995 and aimed ed that more than 50% the St. Lawrence banks at gathering and synthesising much of the data col- between Cornwall and Québec City have lost their lected over the last 30 or so years on species distri- natural integrity, and that shoreline modification has been almost three times greater in this freshwater *See Documents Cited section. section, where human densities are the highest in the nN nN 552 province, than downstream in the estuary and gulf sections of the river (Picard et al. 1997*). Because herpetofaunal populations are sensitive to human disturbance (Green 1997; White et al. 1997), monitoring species diversity is essential to trigger early warnings permitting effective measures to be adopted to deal with threats to ecosystem integrity. This paper presents a regional analysis of the diversi- ty of herpetofauna living along the St. Lawrence River and identifies areas to be protected for the maintenance of amphibian and reptile populations in southern Québec. Study Area and Methods The St. Lawrence Valley Natural History Society, in collaboration with the Société de la faune et des parcs du Québec (FAPAQ), manages a databank containing observations of amphibians and reptiles in Québec gathered by amateur volunteers and pro- fessionals since the end of the 19" century, including historic museum records. Analysis of the data gath- ered prior to 1993 led to the publication of the Atlas of Amphibians and Reptiles of Quebec (Bider and Matte 1996). The data used in our study were extracted from this databank updated in 1996. It con- tains 4010 records obtained along the St. Lawrence River system. More than 55% of these observations were made during the period 1993-1996; i.e.; addi- tional data gathered since the latest edition of the Atlas, and 85% of all observations were made between 1988 and 1996. Common and scientific names of amphibian and reptile species follow Alvo and Oldham (2000). For this project, the study area encompasses the entire St. Lawrence River banks located in Québec from the Québec-Ontario border near Cornwall (45° O01’ N, 74° 43’ W), Ontario, to its northeasternmost area of Blanc-Sablon (51° 25’ N, 57° 08’ W) near the Labrador border. All data collected along the river and within 10 km of its riverbanks, including the Montréal archipelago, the Saguenay River, and the Gulf of St. Lawrence (Baie-des-Chaleurs, Magdalen Islands) were included in the analyses. Ducruc et al. (1995) presented an ecological refer- ence framework developed at the scale of the entire province which is divided into terrestrial units based on stable parameters representing the physical envi- ronment, and characterized with several ecological variables. We present the portrait of the diversity of amphibians and reptiles at the level of the six natural provinces found along the St. Lawrence River, rang- ing from the deciduous forests of the St. Lawrence Lowlands to the tundra vegetation of the Lower North Shore (Figure 1), where the geographic coor- dinates of each record were used to assign them to a natural province. In addition to these natural provinces, records of amphibians and reptiles were also superimposed on THE CANADIAN FIELD-NATURALIST ” | Vol. 116 - an arbitrary division consisting of a systematic grid composed of 100-km? quadrats (10 X 10 km). A mathematical model was used to divide the study area into 751 adjoining quadrats, starting at the southernmost point in Québec (the Québec-Ontario border). All quadrats located less than 10 km from the banks of the St. Lawrence River were included in the study area. Only the presence of a species in a geographic unit, regardless of the number of records and of the species abundance, was used in species richness cal- culations (total number of species in each geographic unit). We acknowledge that species abundance would have provided a more accurate portrait of the regional distribution of amphibians and reptiles but this option was discarded because abundance values were missing from nearly half (44%) of the 4010 records. Global (GRANKs) and provincial (SRANKs) con- servation priority ranking of species, as provided by NatureServe (1999*) and the Centre de données sur le patrimoine naturel du Québec (CDPNQ 1999*), are listed in Alvo and Oldham (2000) and were used to identify species at risk. The Québec priority ranking of S1 was given to species most at risk, compared with S5 for abundant species. A composite prioritization index for the 100-km* quadrats was calculated by attributing 5, 3 or 1 point to species with Québec priority ratings of S1, $2 and S3 respectively, corresponding to the species requiring the most attention from a biodiversity conservation standpoint. The importance of each quadrat in terms of the priority species it contains was assessed with a composite index of prioritization comparing the total points calculated per quadrat with the highest possi- ble total of 31 for a quadrat which would contain all priority species located along the St. Lawrence (S1 = 3 species, S2:= 3 species, $3: = 7 species: Composite index = (n species S1_X_5)+(n species $2 X 3) +(n species $3 X 1) 31 (maximal possible index value) Results Herpetofauna richness The herpetofauna found along the St. Lawrence River is composed of 33 species: 18 amphibians and 15 reptiles. The most commonly observed amphib- ians were the American Toad (Bufo americanus), Spring Peeper (Pseudacris crucifer), Northern Leopard Frog (Rana pipiens) and Wood Frog (Rana sylvatica), while the most abundant reptiles were the Common Garter Snake (Thamnophis sirtalis), Painted Turtle (Chrysemys picta) and Common Snapping Turtle (Chelydra serpentina) (Table 1). Species richness was significantly greater in the natu- ral provinces of southern Québec than in the more easterly ones (Table 1; Figure 1). There were over 32 2002 i Amphibians L1 Reptiles aaa 0 JOBIN, RODRIGUE, AND DESGRANGES: AMPHIBIAN AND REPTILE DIVERSITY 100 | % on ro) Lowlands Appalachians Laurentians 553 A: Lowlands B: Appalachians C: Laurentians D: Lower North Shore Plateau E: Anticosti Island F: Magdalen Islands Cc QUEBEC NEWFOUNDLAND AND LABRADOR : NEW PEA BRUNSWICK MAINE (USA) NOVA SCOTIA Lower North Shore Plateau Anticosti Island Magdalen Island FIGURE 1. Number and percentage of amphibian and reptile species observed in the six natural provinces along the St. Lawrence River in Québec. Inset shows the localisation of natural provinces and their 10km-wide zone along the St.Lawrence River where amphibian and reptile records were compiled. species in the St. Lawrence Lowlands versus fewer than ten species on the Lower North Shore Plateau, Anticosti Island and the Magdalen Islands. While the number of reptile species decreases as one moves downstream along the St. Lawrence River, amphibian diversity remains fairly steady throughout the fresh- water section and estuary and then rapidly declines in the Gulf. Although found elsewhere in Québec, three species (Spring Salamander [Gyrinophilus por- phyriticus], Mountain Dusky Salamander [Desmognathus ocrophaeus|, Common Musk Turtle [Sternotherus odoratus]) and one subspecies (Boreal Chorus Frog [Pseudacris triseriata maculata] were not recorded along the St. Lawrence River. The richest sites for both amphibians and reptiles are located in southwestern Québec around the Lac des Deux-Montagnes, Lac Saint-Louis and Lac Saint-Frangois. The richest quadrat in the river sys- tem was the [le Perrot quadrat where 16 species of amphibians and 13 species of reptiles were recorded, followed by the quadrats along the Lac des Deux- Montagnes (16 amphibians, 9 reptiles) and in the Senneville—Sainte-Anne-de-Bellevue area (12 amphibians, 8 reptiles). Quadrats in the Valleyfield (12 amphibians, 7 reptiles) and Saint-Clet regions (13 amphibians, 5 reptiles) also showed high species richness. The Baie de Beauport quadrat in the Québec City region is noteworthy among the quadrats east of Montréal for its high number of species (12 amphibians, 6 reptiles). The Magdalen Islands had only one record for a reptile (one Leatherback [Dermochelys coriacea]), while Anticosti Island had none. Maps showing all of the 100-km? quadrats where each species was observed along the St. Lawrence River are available on The Biodiversity Portrait of the St. Lawrence web site (DesGranges and Ducruc 2000*). Species of concern Four amphibian and nine reptile species observed along the St. Lawrence River have a high-priority conservation status in Québec (Table 1) but their populations would not be threatened globally (global ranks = G4 or G5). In addition, the Leatherback has a high-priority status globally (G3) but was not con- sidered threatened in Québec (SZN) due to its spo- radic occurence along its coasts. Most records for species of concern came from the Montréal region, where 11 out of 13 listed species were observed around the Lac des Deux-Montagnes and Lac Saint- Louis (Figure 2). The quadrats with the highest pri- oritization index (Table 2) were also those with the highest species richness (Ile Perrot, Pointe-Claire, Lac des Deux-Montagnes, Senneville). Several neighbouring quadrats (Roxboro, Laval, Laval-north, Mont-Royal) also supported many species of con- cern. Some observations of species of concern were made east of Lac Saint-Pierre such as the Wood Vol. 116 10 10 10 V0 c0 © 0 v0 9°0 L’‘0 al 81 ~ 36% re Uv Ee) c0 v0 L0 i! 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JOBIN, RODRIGUE, AND DESGRANGES: AMPHIBIAN AND REPTILE DIVERSITY 555 St.Lawrence River MONTREAL 1 Mont-Royal = ia yh bins’ : Lac Saint-Francois LYS ast a w<>E inte- ne ointe-Clair (Pointe-Fraser) we ei aire s ore hed lle Perrot 0 25 50km Ficure 2. Value of the composite index of prioritization of herpetofauna for quadrats located in the Montréal region. Turtle (Clemmys insculpta) and Brown Snake (Storeria dekayi) recorded in a few quadrats in the Québec City region and in the Laurentians and Appalachians provinces. Discussion Unlike previous publications on amphibian and reptile species distribution in eastern Canada where observation records were either mapped (Bleakney 1958; Logier and Toner 1961) or analysed to draw generalized distribution maps for each species (Cook 1984), we present a synthetic view of species rich- ness for distinct geographic regions without specific emphasis on the actual distribution of each species. Our analysis clearly showed that amphibian and rep- tile species richness decreases as one moves down- stream along the St. Lawrence River from the Montréal region to the Lower North Shore and the Gulf. Interestingly, the region of the Montréal archipelago showing the highest species richness corresponds well with Bleakney’s (1958) herpeto- faunal section 2 considered as the richest region in Québec which extends along the Outaouais River Valley, in the St.Lawrence Lowlands from the Québec-Ontario border down to Lake Saint-Pierre and extending over the Richelieu River drainage basin. Several factors could contribute to a lower her- petofaunal species richness in the northeastern regions. Climatic conditions are harsher in the cold regions of the Lower North Shore Plateau and Anticosti Island which may limit species range (Bleakney 1958; Bider and Matte 1996; McKenney et al. 1998). Several species, including the Four-toed Salamander (Hemidactylium scutatum), Common Map Turtle (Graptemys geographica) and Spiny Softshell (Apalone spinifera), are at the northern limit of their range in southwestern Québec, their absence in colder regions thus contributing in a lower species richness beyond the St. Lawrence Lowlands. The lower diversity of habitats in the northeastern regions as compared to southern regions would also limit species distribution and species associated with specific habitat types should only be found in regions where this habitat occurs. For example, the Western Chorus Frog (Pseudacris tris- eriata) and the Smooth Green Snake (Liochlorophis [Opheodrys] vernalis) are generally associated with open fields (Cook 1984; Bider and Matte 1996) which are uncommon in the gulf regions thus increasing species richness in the south. Amphibian species observed on the Magdalen Islands and Anticosti Island were probably introduced, with the exception of the Mink Frog (Rana septentrionalis), which may be native to Anticosti Island (Bider and Matte 1996). 556 THE CANADIAN FIELD-NATURALIST Vol. 116 TABLE 2. Quadrats with the highest composite index of prioritization Cc Species observed with conservation rank of omposite Quadrat! Index Sl S2, SS Ile Perrot 0.81 Spiny Softshell Blanding’s Turtle Four-toed Salamander Common Map Turtle Wood Turtle Western Chorus Frog Brown Snake Pickerel Frog Ringneck Snake Milk Snake Northern Water Snake Pointe-Claire 0.39 Common Map Turtle Blanding’s Turtle Milk Snake Brown Snake Roxboro 0.39 Common Map Turtle Wood Turtle Milk Snake Brown Snake Lac des Deux-Montagnes 0.39 Common Map Turtle Brown Snake Western Chorus Frog Pickerel Frog Ringneck Snake Northern Water Snake Senneville 0.35 Common Map Turtle Brown Snake Western Chorus Frog Milk Snake Northern Water Snake Laval O32 Spotted Turtle Common Map Turtle Laval (north) 0.29 Common Map Turtle Blanding’s Turtle Milk Snake Mont-Royal 0:29 Common Map Turtle Brown Snake Milk Snake Lac Saint-Frangois 0.23 Common Map Turtle Pickerel Frog (Pointe-Fraser) Northern Water Snake See Figure 2 for quadrat location Differential searching effort among regions would likely explain part of the variation of the observed species richness. Because Québec’s human densities are at their highest in southern Québec, volunteers are also more numerous in this part of the province. This is reflected in the higher number of records coming from southern Québec. Because our analysis relied on records gathered over decades by volun- teers and professionals under varied scenarios and using different methodologies, there were no consis- tent and rigorous procedures followed in data acqui- sition. It is therefore difficult to evaluate the relative importance of many factors (climate, habitat, search- ing effort) on the actual species distribution patterns. Our analysis, however, presents the best portrait of amphibian and reptile species distribution along the St. Lawrence based on the most up-to-date informa- tion available in Québec. Initial distribution maps of Canadian amphibian and reptile species were published in Bleakney (1958), Logier and Toner (1961) and Cook (1984), and based largely on verifiable museum specimens. Comparison of these maps with records of species from the St. Lawrence River databanks based largely on reported observations reveals numerous addition- al localities gathered, especially since the Atlas of Amphibians and Reptiles in Québec project has been instigated, which have extended the presumed distri- bution of several species further north and east of their previously known limits (see Bider and Matte 1996). New locations mapped in Bider and Matte (1996) of the Four-toed Salamander and Spotted Turtle (Clemmys guttata) in the St. Lawrence Lowlands, the Gray Treefrog (Hyla versicolor) in Lake Saint-Pierre, the Common Snapping Turtle east of Québec City, and the Bullfrog (Rana catesbeiana), Spotted Salamander (Ambystoma maculatum) and Redbelly Snake (Storeria occipitomaculata) in Lake Saint-Jean are notewor- thy. Most interestingly, additional locations of the Blanding’s Turtle (Emydoidea blandingii) in the Montréal region and in the freshwater estuary sec- tion of the St. Lawrence west of Québec City have been reported to the data bank since the latest edition of the Atlas of Bider and Matte (1996). Caution must be exercised, however, in accepting observations 2002 where no voucher specimens or photographs are available for verification. As well, all turtle records which “extend” a known range are initially most sus- pect because of the likelihood that they are based on escaped captives brought from natural populations elsewhere. Despite such reservations, deletion of suspect records would not significantly alter the gen- eral pattern described here. Species of concern Species of high-priority conservation status in Québec are generally designated as such because they are at the northern limit of their range, are uncommon, and are sensitive to habitat destruction (Beaulieu 1992; Bider and Matte 1996). However, with the exception of the Leatherback, global status populations are generally not of concern. Among the species with the highest conservation status in Québec (S1) which are observed along the St. Lawrence River, two (Spiny Softshell, Spotted Turtle) are included in a previous list of rare and endangered Canadian amphibian and reptile species (Cook 1970), but the Common Map Turtle is lack- ing from this list. Several other species of concern in Québec such as the Wood Turtle (S2), Brown Snake (S2), Four-toed Salamander (S3), Western Chorus Frog (S3), Pickerel Frog (Rana palustris; S3), Northern Water Snake (Nerodia sipedon; S3), Milk Snake (Lampropeltis triangulum; S3), and Ringneck Snake (Diadophis punctatus; S3) were also absent from Cook’s list which was based on total Canadian ranges. As well, these discrepancies in status attribution result from increased knowledge of species distribution and numbers acquired since Cook’s list was published and from more objective criteria now applied by the CDPNQ for the province of Québec. Eleven out of the 13 species with a high-priority conservation status in Québec were observed around the Lac des Deux-Montagnes and Lac Saint-Louis in the Montréal region making this a high-priority area for the conservation of the diversity of Québec’s her- petofauna. Because these species contribute to the biodiversity of the province of Québec, conservation efforts should be directed at maintaining their local populations. In addition, the Montréal archipelago has been identified as a major sector for biodiversity conservation in Québec based on local species rich- ness and priority index calculated for major taxo- nomic groups (fish, vascular plants, herpetofauna, breeding birds) (DesGranges 2000*). Although sev- eral areas are currently protected in the Montréal region, land managers need to be aware of anthro- pogenic pressures (e.g., pollution, habitat loss, fluc- tuating water levels) being exerted on remaining high-quality habitats not currently subjected to any conservation status and should act accordingly through land acquisition or stewarship programs in order to protect these communities. JOBIN, RODRIGUE, AND DESGRANGES: AMPHIBIAN AND REPTILE DIVERSITY Se | Acknowledgments We are indebted to the more than 350 volunteers across the province of Québec whose observations still continue to enrich the databank used in this study. We thank Jacques Jutras of the Société de la faune et des parcs du Québec (FAPAQ) for his involvement in the Atlas program and the financial support of the FAPAQ, and J. Roger Bider who initi- ated the project leading to the production of the Atlas of the amphibians and reptiles of Québec. We also thank Luci Bossé who kindly provided informa- tion on the Leatherback Turtle distribution in the Gulf of St. Lawrence. Documents Cited (marked * in text) CDPNQ (Centre de données sur le patrimoine naturel du Québec). 1999. Centre de données sur le patri- moine naturel du Québec (CDPNQ). Société de la Faune et des Parcs du Québec, Gouvernement du Qué- bec, Québec. DesGranges, J.-L. 2000. Protecting the biodiversity of the St. Lawrence: Conservation plan. Jn Biodiversity Portrait of the St. Lawrence. Edited by J-L. Des Granges and J.-P. Ducruc. Canadian Wildlife Service, Environment Canada, Québec Region and the Direction du patrimoine écologique, Ministére de 1’ Environ- nement du Québec. (www.qc.ec.gc.ca/faune/biodiv). DesGranges, J.-L., and J.-P. Ducruc. Editors. 2000. Biodiversity Portrait of the St. Lawrence. Canadian Wildlife Service, Environment Canada, Québec Region and the Direction du patrimoine écologique, Ministére de l1’Environnement du Québec. (www.qc.ec.gc.ca/ faune/biodiv). NatureServe. 1999. NatureServe Central Databases. Arlington, Virginia. U.S.A. (Retrieved March 1999). Picard, M., D. Lehoux, R. Langevin, and C. Grenier. 1997. Etat des rives et protection des milieux humides du fleuve Saint-Laurent: Synthése des connaissances actuelles. Environnement Canada, Service canadien de la faune, Région du Québec. 68 pages. Literature Cited Alvo, R., and M. J. Oldham. 2000. A review of the status of Canada’s amphibian and reptile species: a compari- son of three ranking systems. Canadian Field-Naturalist 114: 520-540. Beaulieu, H. 1992. Liste des espéces de la faune vertébrée susceptibles d’étre désignées menacées ou vulnérables. Gouvernement du Québec, Ministére du Loisir, de la Chasse et de la Péche. 107 pages. Bider, J. R., and S. Matte. 1996. The Atlas of Amphibi- ans and Reptiles of Quebec. St. Lawrence Valley Natural History Society and Ministére de |’ Environne- ment et de la Faune, Direction de la faune et des habi- tats, Québec. 106 pages. Bishop, C. A., and K. E. Pettit. Editors. 1992. Declines in Canadian amphibian populations: Designing a national monitoring strategy. Canadian Wildlife Ser- vice, Ottawa, Occasional Paper Series (76). 120 pages. Blaustein, A. R., and D. B. Wake. 1990. Declining amphibian populations: A global phenomenon? Trends in Ecology and Evolution 5: 203-204. 558 Bleakney, J. S. 1958. A zoogeographical study of the amphibians and reptiles of eastern Canada. National Museum of Canada, Bulletin Number 155, Biological Series Number 54, Ottawa, Ontario. 119 pages. Cook, F. R. 1970. Rare or endangered canadian amphib- ians and reptiles. Canadian Field-Naturalist 84: 9-16. Cook, F. R. 1984. Introduction to Canadian Amphibians and Reptiles. National Museums of Canada, Ottawa. 200 pages. Ducruc, J.-P., T. Li, and J. Bissonnette. 1995. Small- scale ecological mapping of Québec: Natural provinces and regions (cartographic delineation). Pages 45-53 in Landscape Ecology in Land Use Planning: Methods and Practice. Edited by G. Domon and J. Falardeau. Proceedings of the fourth workshop of the Canadian Society for Landscape Ecology and Management, Université Laval, Sainte-Foy, Québec, June 1994. Polyscience Publications Inc., Morin Heights, Québec. 227 pages. Green, D. M. Editor. 1997. Amphibians in decline: Canadian studies of a global problem. Herpetological THE CANADIAN FIELD-NATURALIST Vol. 116 - Conservation (1), Society for the Study of Amphibians and Reptiles. Saint Louis, USA. 338 pages. Logier, E. B. S., and G. C. Toner. 1961. Checklist of the amphibians and reptiles of Canada and Alaska. Second Edition. Royal Ontario Museum, Toronto, Ontario. 92 pages. McKenney, D. W., B. G. Mackey, J. P. Bogart, J. E. McKee, M. J. Oldham, and A. Chek. 1998. Bioclimatic and spatial analysis of Ontario reptiles and amphibians. Ecoscience 5: 18-30. Phillips, K. 1990. Where have all the frogs and toads gone? A recent workshop described an apparent decline worldwide of amphibian populations. BioScience 40: 422-424. White, D., P. G. Minotti, M. J. Barezak, J. C. Sifneos, K. E. Freemark, M. V. Santelmann, C. F. Steinitz, A. R. Kiester, and E. M. Preston. 1997. Assessing risks to biodiversity from future landscape changes. Conservation Biology 11: 349-360. Received 28 December 2000 Accepted 22 November 2002 First Report of the Rare Charophyte Nitella macounii (T. F. Allen) T. F. Allen in Saskatchewan and Western Canada HENRY MANN! and M. V. S. Rasu2 !Environmental Science/Biology, Sir Wilfred Grenfell College, Memorial University of Newfoundland, Corner Brook, Newfoundland A2H 6P9 Canada 2Biology Department, University of Regina, Regina, Saskatchewan S4S 0A2 Canada Mann, Henry, and M. V.S. Raju. 2002. First report of the rare charophyte Nitella macounii (T. F. Allen) T. F. Allen in Saskatchewan and western Canada. Canadian Field-Naturalist 116(4): 559-570. Because few collections of Nitella macounii exist and early descriptions were largely based on dried herbarium material, a number of morphological features such as type and form of branching have been reinterpreted suggesting reclassifica- tion into Section Palia from Section Nitella. Earlier observations that the species may be related to N. stuartii are support- ed and a potential disjunct range with Argentina seems a possibility. Oospore membrane decoration is described in detail and conflicting earlier reports clarified. Especially significant is the first report that this species is tetrascutate, only the sixth charophyte known to be so. The ephemeral habitat and the ecology of N. macounii in Saskatchewan are discussed in light of the recent charophytivory hypothesis which predicts that such species are restricted to habitats where aquatic invertebrate herbivores are absent or present in low numbers. The dynamic nature of ephemeral habitats both in time and space is postulated to be a prime factor in the distribution and rarity of this species throughout its range. It is suggested that because of its unexpected ephemeral ecology, N. macounii may be more widespread across the Great Plains of North America than is presently known. Currently it must be considered globally and nationally very rare with only 16 known collection sites. Key Words: Charophytes, Characeae, Nitella macounii, Nitella stuartii, Chara braunii, Saskatchewan, ephemeral prairie wetlands. The charophytes of Saskatchewan and the adjacent provinces are poorly documented. Only one rather restricted survey has been carried out (Mann 1994b) confirming that few collections from the area exist and almost no descriptive or distributional literature is available. The region is rich in varied aquatic habitats, including the oligotrophic waters of the northern bore- al forest, the eutrophic lakes, streams and sloughs of the prairies, saline lakes of varying sizes that dot the south, and a great variety of ephemeral and semi- permanent wetlands on the cultivated plains and the cattle ranges (Warner and Rubec 1997; National Wetlands Working Group 1988). Because a variety of habitats tend to engender a variety of species, each with its own particular adaptations to the local biotic and abiotic conditions, it should be anticipated that much taxonomic, ecological and biogeographic charo- phyte data will be collected from this region if and when intensive investigations are undertaken. That an aquatic charophyte flora exists in the often dry and only intermittently shallow flooded ephemeral sites of intensively cultivated grain fields is perhaps a surprise to some, but it is consistent with the broad range of ecological adaptations known for charophytes as well as recent studies such as those by Proctor (1990; 1999) from the southern extremity of the Great Plains. Our observations suggest that charophytes are com- mon and ubiquitous in most Saskatchewan aquatic habitats and that each set of conditions fosters its own particular assemblage of species. Here we report on several unique aspects of prairie charophytology. This is the first report of the rare charophyte Macoun’s Nitella (Nitella macounii (T. F. Allen) T. F. Allen) from Saskatche- wan and from Canada west of the Great Lakes. Features of this new distribution suggest that an expanded range may exist on the Great Plains for this species as well as a possible disjunct range with South America. Secondly, little is known about the detailed ecology of most charophyte species. We here present and discuss a somewhat novel ecological situation along with some conjec- tures which may lead to future investigations. Thirdly, the Saskatchewan specimens are compared to the literature and to historical collections and several issues such as oospore membrane ornamen- tation and antheridial structure are clarified. It is especially important to redescribe some of the sig- nificant morphological features of this species since few specimens exist and many of the details described in the literature have been derived from only a small number of plants, mostly dried herbar- ium specimens from which some morphological detail is difficult to interpret. Sufficient live and/or liquid preserved specimens are necessary to fully appreciate the gross morphology and range of vari- ation normal for a species. New and re-interpreted morphological features for N. macounii point to a taxonomic re-alignment from the traditional view of Wood (1965). 59 560 Study Site and Methods The core area in which N. macounii was discov- ered and intensively sought is centered in the village of Belle Plaine, Saskatchewan, located midway between the cities of Regina and Moose Jaw on the Trans-Canada Highway (Canadian Topographic Map, Drinkwater, 72-I/6, Edition 4, 1988, Grid Zone Designation 13U, Square Identification DF, Energy, Mines and Resources Canada). Belle Plaine lies within Landscape Area 17 known as the Regina Plains and within the greater ecoregion designated as Moist Mixed Grassland (Fung 1999). The area is a flat lacustrine plain with dark brown clayey soils and is almost totally cultivated. Few permanent water bodies exist in the study area. Infrequent small sloughs hold water throughout the summer in most years as do farm dugouts and road building borrow pits. Shallow depressions accumulate meltwater in spring, but by late spring to mid-summer these are frequently dry and often cultivated. Of the average annual total precipitation (380 mm), most falls dur- ing the summer months in seemingly erratic patterns over the landscape. Thunderstorms are common dur- ing the summer, often dumping considerable rain in localized areas. The apparent randomness of thun- derstorm precipitation results in a patchwork of drier and wetter localized sites within the larger climatic fluctuation patterns of drier and wetter years. Areas which receive a number of spaced downpours during the season may have considerable standing water in shallow ephemeral depressions throughout the grow- ing season, while adjacent areas may be much drier with little or no standing water (Euliss et al. 1999). As such, habitat for species adapted to ephemeral aquatic sites will exist in a dynamic flux over the landscape from season to season. Surveys encompassed a radius of about 10 km around Belle Plaine. Four collections were made, three in 1996 and one in 2000. Site 1 (tractor rut) of 15 July 1966 has a map grid reference of 894E, 823 N. Site 2 of 18 July 1996 was within 200 meters of site 1 (an inundated fallow farm field). Site 3 of 18 July 1996 (inundated stubble farm field) has a grid reference of 882E, 823N. Site 4 of 25 July 2000 is situated in a temporary marsh/farm field with a grid reference of 872E, 729N. Visits were made to sites 1, 2, and 3 in the summers of 1997, 1998, and 1999, but they were dry early in the season and no charo- phytes were located. Surveys were also made in 1996 and 2000 to the north-east of Regina, east to Indian Head and the Strawberry Lakes area, as well as in the Grayson, Crooked Lake, Melville, and Yorkton areas. Charophytes were collected by hand and either preserved in 5 percent formalin or dried as herbari- um specimens. All critical examinations and mea- surements were made on liquid preserved material. One hundred measurements were made of each THE CANADIAN FIELD-NATURALIST Vol. 116 - gametangial feature distributed approximately equal- ly among at least four different individuals. Mea- surements and illustration are consistent with procedures described in Mann (1994b). Spore prepa- ration for the SEM (Scanning Electron Microscope) followed the method of John and Moore (1987). Material from site 4 for chromosome counts was fixed in Farmer’s Fluid and stained with the Iron Alum/Acetocarmine method of Godward (1948). Voucher specimens of N. macounii are housed in the Sir Wilfred Grenfell College Herbarium (SWGC) and duplicates have been deposited with the Phycological Herbarium of the Canadian Museum of Nature, Ottawa (CANA) and with the Newfoundland Museum, St. John’s (NFM). As well, Saskatchewan specimens of Chara braunii have been deposited with the above institutions. Herbaria abbreviations used are listed in Holmgren et al. (1990) except for SWGC which is not yet listed. The nomenclature of Looman and Best (1987) is used for vascular plants and Clifford (1991) for invertebrates. The following specimens were examined for com- parative purposes on loan from their respective herbaria: (a) Jas. M. Macoun, August 15, 1884, St. Clair River, Ontario (CANA 5840). (b) Allen and Saunders, No. 4010, August 1898, Clear Lake, South Dakota (FH). (c) Glenn Crum, No. 499, July 2, 1974, Wright County, Iowa. Sheet 1 of 3 and sheet 3 of 3 CISC 424538 and 424539). Morphology Saskatchewan N. macounii is a small plant up to 11 cm in height arising as a single axis in dendroid fashion from a basal rhizoidal node, or in a fruticose fashion with several axes arising from the basal node. Main axes are slender and up to 600 wm in diameter. In the 2000 collection plants growing in only a few centimeters of water were very tiny, 3.0 to 3.5 cm in height. Plants are superficially very Tolypella-like with apparent long sterile branchlets at the lower nodes, usually more than two branches at a node, and forming small dense heads (Figure 1A). T. F. Allen (1887) indicates the species has “fertile heads” and “imperfectly formed nests”, as do G. O. Allen (1954) and Crum (1975). Wood (1965) states that “heads are not formed, the entire upper parts resembling a head.” Wood and Imahori (1964) state that mucus is absent, but Saskatchewan plants exhibit some mucus within the small compact heads. This feature is not mentioned elsewhere in the litera- ture for this species; however, it is well known that small amounts of mucus are difficult to detect in dried herbarium specimens. The ability of a species to produce mucus has been used as a taxonomic - character by most authors and there is some indica- tion that it may be a useful feature in determining evolutionary radiation and dispersal in the Nitelleae (Proctor 1997). 2002 MANN AND RAJU: THE CHAROPHYTE NITELLA MACOUNII 561 FiGuURE 1.A. Individual plant of N. macounii. Abbreviations: tc, termi- nal cluster; bn, basal rhizoidal node; fn, first node; sb, long “ster- ile” branchlet at first node. Scale bar = 1.0 centimeter. B. Terminal cluster composed of several dense heads. Scale bar = 0.25 centimeter. C. Individual “prickly” dense head. Scale bar = 0.25 centimeter. D. Individual larger branchlet of dense head which produces the protruding dactyls creating the “prickly” appearance. Scale bar = 0.15 centimeter. 562 At the first node above the basal rhizoidal node long “sterile” branchlets occur, up to 6.5 cm in length, sometimes simple and non-furcate with sev- eral (1-3) shorter end cells, and sometimes once or twice (rarely thrice) furcate. Originally T. F. Allen (1888) and also Wood (1948) considered this species to be heterophyllous (= heteroclemous; 1.e., having branchlets of two types in a whorl); however, it is now considered to be homoeophyllous (= homoeo- clemous), at most, dimorphic (G. O. Allen 1954). Some Nitellas such as N. hyalina and N. stuartii exhibit distinctly different branchlet forms (acces- sory branchlets) in their whorls (1.e., are hetero- clemous) and this feature has been used taxonomi- cally for identification purposes. In our opinion, N. macounii shows a heteroclemous tendency, especial- ly in lower whorls, but-it is not at all clear or well defined. G. O. Allen (1954) states, and present authors concur, that considerable abscission and/or sup- pressed furcation occurs giving the appearance of deviation from the basic 2-furcate branchlet condi- tion and producing the appearance of multicelled dactyls. Beyond the first axis node the 2-furcate, sin- gle-celled dactyl condition becomes more regular with increasing nodes. The literature (Crum 1975; Wood 1965) indicates 6—8 branchlets per node, but in the Saskatchewan material sometimes up to 12 branchlets have been noted at the first node and from 6—8—10 at subsequent nodes. Whether some of these are in fact accessory branchlets is not clear. Beyond the first node branchlets become shorter, more regu- larly 2-furcate (occasionally 3-furcate) and become fertile with clusters of 2-5 oogonia at the first and 1-3 oogonia at the second furcations, and a single antheridium terminating each furcation. Even a long “sterile” branchlet at the first node may very occa- sionally exhibit an oogonium. Antheridia appear to abscise early and can be found only on the smaller branchlets and those of the dense heads. Branchlets generally have 2—5 secondary rays, each ray bearing 2-4, occasionally 5, one-celled acute-acuminate dactyls. Dactyls of the dense heads tend to be more acuminate (Figure 1D). Axes exhibit up to 5 nodes. Four to six branches occur at each node, the branch axis internodes becoming progressively shorter to form terminal clusters (Figure 1B) composed of smaller clusters, which themselves are composed of the tightly packed dense heads having a prickly appearance (Figure 1C). The dense center of these heads, ignor- ing the longer projecting dactyl tips, range from 0.1 to 0.5 cm in diameter. Dactyls of the cluster branch- lets and outer dactyls of the dense heads are moder- ately encrusted with the sharp points of the acute/acuminate tips protruding leading one to won- der whether this “porcupine” condition of the prickly dense heads provides a mechanical deterrent to small invertebrate herbivores, perhaps especially relevant THE CANADIAN FIELD-NATURALIST | 4 : Vol. 116 - to the delicate antheridia which are mainly found in the tighter clusters and in the dense heads. Gametangial measurements are given in Table 1 and compared to other reports for the species. Only Wood (1965) provides a full range of measurements and these are apparently taken from only two speci- mens, the Niagara holotype and a Wilmot, South Dakota specimen of T. F. Allen, both dried herbari- um material. G. O. Allen (1954), after examining Lake St. Clair material, suggests that T. F. Allen’s (1887; 1894) oospore and antheridial measurements are too low. The measurements of Crum (1975) are the largest reported. We have examined Crum’s col- lection No. 499, July 2, 1974 which are vigorous, freely fruiting plants up to 12 cm tall. Although there is some indication that ecological factors are capable of modifying gametangial size (Griffin 1963), the extent to which this is possible is not known for the different charophyte species. The moderate differ- ences seen in Table | may be considered to fall with- in the normal genetic variation range of the species. At maturity the small antheridia of N. macounii, are greenish-yellow in live material and are tetrascu- tate. Most charophytes are octoscutate. No previous accounts of this species have observed this antheridi- al feature. Only the following five charophytes in addition to N. macounii are known to be tetrascutate: Chara zeylanica Klein ex Willd. (Proctor et al. 1971), N. terrestris lyengar (Iyengar 1958), N. stuar- tii A. Braun (Carl de Donterberg and Rotman 1973), N. cordobensis Caceres (Caceres 1975), and N. quadriscutulum Jao and Li (Han and Li 1994). Tuttle (1924) also mentioned a tetrascutate charophyte from Virginia but it is not possible to verify this or determine the taxon. Because the tetrascutate condi- tion is difficult to determine from dried herbarium specimens, especially in species with tiny antheridia whose plates do not separate easily, this feature may be more common in charophytes than is presently known. Most species descriptions in the literature make no reference to the number of antheridial scutes. The tetrascutate feature has been shown to be taxonomically significant in the elucidation of the Chara zeylanica complex (Proctor et al. 1971) and may also prove to be so in the genus Nitella. Oospore membrane decoration has long been con- sidered an important taxonomic feature in the genus Nitella (Groves and Bullock-Webster 1920). More recent studies with the SEM have further elucidated details of membrane ornamentation in selected species (Frame 1977; John and Moore 1987; Casanova 1991). The strongly ornamented oospore of N. macounii is very characteristic of the species, yet considerable discrepancy exists in the literature. T. F. Allen originally described it as smooth due to the lack of mature spores in the original collection, but later upon receiving material with mature oospores described it as having a surface marked by short elevations like mountain peaks, somewhat 2002 MANN AND RAJU: THE CHAROPHYTE NITELLA MACOUNII 563 TABLE 1. Gametangial and oospore features of Nitella macounii. Measurements are presented as means (um) with range values in parentheses. This study a. F Allen G. O. Allen Wood Crum 2002 1887 & 1894 1954 1965 1975 Oogonium length* 386 (341-413) - (345-—360)* (350-450) width 283 (248-310) ~ ~ (255-270) (320-350) coronula height 44 (36-52) sa x (42-53) at coronula width 65 (52-72) - a (60-80) a convolutions (6-8) = == (7-8) r Oospore length 243 (207-248) (210-215) 275 (225-240) (260-280) width 241 (207-248) (200-210) 250 (180-225) (240-260) No. of ridges (4-6) (5-8) (5-6) (5-8) (5-6) fossae width 53 (41-62) — = 42 - Antheridium diameter 191 (165-227) 130 ~ (195-225) (200-250) * All measurements include coronula height except Wood (1965) which does not. elongate and arranged irregularly at right angles over the fossae surfaces (T. F. Allen 1894). Wood (1948) describes it in the same fashion, but later (Wood 1965) states it is “strongly vermiferous, the irregular raised areas occasionally becoming imperfectly reticulate.” G. O. Allen (1954) does not consider T. F. Allen’s (1894) description and illustration to be accurate, but describes it as a “coralloid reticulum, i.e. a deeply sculptured but imperfect reticulum, looking rather as if some fish-netting had gone rotten and became broken in many places.” Crum (1975) simply states the membrane to be strongly discontin- uous reticulate. The only SEM view of this species (Frame 1977) concludes the membrane to be papil- late-reticulate to strongly reticulate when mature. The present study demonstrates that all of the above views on the oospore membrane decoration of N. macounii have some validity, but they each only address a part of the total picture presumably due largely to a small number of plants and oospores examined. The present study has had available a considerable number of different plants collected from four localities and liquid preserved. Large num- bers of oospores which had washed free of their plants and which had been naturally “released” from their enveloping cells could be and were examined. It became quickly apparent that all naturally shed oospores were not alike. All appeared fully formed and apparently viable, but existed in three distinct ranges as seen with reflected stereoscopic light: pale golden-yellow, pale orange-brown, and dark orange- brown. Quantitatively, these were approximately present as 10 percent, 20 percent, and 70 percent respectively. Literature reports for oospore color include dark brown (Wood 1965), dark chestnut brown (Wood and Imahari 1964), and deep golden- brown (Crum 1975). The pale golden-yellow spores exhibit a distinct fine reticulum with approximately 9-11 reticulation meshes across a fossa (Figure 2D). At the other extreme, the darkest orange-brown spores exhibit a distinct tuberculate series of irregu- lar raised areas as described and illustrated by T. F. Allen (1894), Wood and Imahori (1964), and Wood (1965). This pattern is clearly seen in the SEM Figure 2A, B. Focusing slightly down with the light microscope, near the base of the flaring tubercles, will produce a coarse reticulate view. Casanova (1991) also noted this discrepancy between SEM and light microscope views. The intermediately-colored pale orange-brown spores have an appearance some- what intermediate between the fine reticulation and the coarse tuberculate which might be described as an imperfect “rotten fish-net” reticulum (Figure 2C). Specimens examined on loan (CANA, FH, and ISC) exhibited similar membrane features as those described here. At certain stages N. macounii mem- brane decoration resembles that of Casanova’s (1991) N. stuartii, but even more resembles her tuberculate spores of N. pseudoflabellata. It seems clear that more studies like the present and Casanova (1991) showing the range of ornamentation produced by a taxon are necessary if we are to fully understand the taxonomic significance and limitations of this feature. In the traditional view, the pale golden-yellow spores represent immature spores and the darkest those that are fully mature. All studies are based on this assumption which has not been critically tested. Since all of these spore types of N. macounii are nat- urally shed, they might be considered equally mature, or are immature spores at various stages of development also naturally shed? Can the ornamen- tation of spores further develop after they have been shed? Or is it possible that in this species and others fully mature polymorphic spores are produced, genetically determined, as would be analogous to well known examples of seed polymorphy in some of the angiosperms (Harper et al 1970)? The litera- 564 ture seems to be blank on these questions. It may be suggested that having the ability to produce a variety of spore types with varying wall thicknesses related to length of dormancy would be ecologically advan- tageous to a species utilizing ephemeral habitats which shift spatially and temporally with the vagaries of the climate and seasons. Casanova and Brock (1996) describe timing variation (i.e., poly- physiologic spores) in spore germination of species adapted to ephemeral sites. There seems to be no reason why this feature may not also exhibit a mor- phological expression, at least in some charophytes. In the absence of conclusive evidence, we can only speculate on this matter at present. Taxonomy Nitella macounii (T. F. Allen) T. F. Allen was first described by Allen from two specimens sent to him by J. M. Macoun collected at Niagara Falls in 1882 (T. F. Allen 1887). Because of its Tolypella-like habit, Allen originally named it Tolypella macounii, stating “This is one of the most remarkable Tolypellae that has yet been described .... It is Nitella-like in its habit of growth, slightly encrusted, and altogether unique.” G. O. Allen (1954) examined one of the original Niagara type locality specimens and could find no “ripe fruit” which due to its flat- tened structure would have indicated a Nitella rather than a Tolypella. After receiving additional speci- mens which clearly showed terminal antheridia col- lected by Macoun from Lake St. Clair, T. F. Allen renamed the plant Nitella macounii (T. F. Allen 1888). Allen again published a brief description of the species in his Characeae of America series (T. F. Allen 1894). Subsequent descriptions by R. D. Wood (1948; 1965) utilized earlier descriptions and the examination of one and two specimen collections respectively. G. O. Allen’s Annotated Key (1954) provides some useful observations on branchlet structure and oospore membrane decoration. The most recent brief description is from Iowa by Crum (1975). Wood (1965) includes N. macounii in Subgenus Nitella characterized by one-celled dactyls, and in Section Nitella with non-acuminate dactyl apices. Later in his description of N. macounii Wood contra- dicts this distinction by stating that dactyl apices are acuminate. Included in Section Nitella is Wood’s N. flexilis group and N. mirabilis, both producing one- furcate branchlets, whereas N. macounii has 2- furcate branchlets. Tindall (1967) in describing a new species of Nitella (N. hotchkissii) from the southwestern USA, suggests two similar species groups, the N. flexilis group (including WN. flexilis, N. opaca, N. missourien- sis, N. mirabilis, N. hotchkissii) and the N. acumina- ta group (including N. acuminata, N. stuartii, N. allenii, N. macounii). Both groups show parallel morphological features and each shows a distinct THE CANADIAN FIELD-NATURALIST a EEE Oh ay Aye !!!!!™C*~C 10%) of northern Great Plains land- scapes. Key Words: Cooper’s Hawk, Accipiter cooperii, nesting density, reproductive success, North Dakota, Great Plains The mostly treeless Great Plains of North America provide the primary breeding habitat for many grass- land-dependent bird species (Knopf 1996), but little habitat for many woodland species such as Accipiter hawks. For example, breeding Cooper’s Hawks (Accipiter cooperii) have been thought to be rare in the western Dakotas (Stewart 1975:90; South Dakota Ornithologists Union 1991:74) and in southeastern Alberta (Semenchuk 1992:83). Indeed, this hawk has been regarded as a species of special concern in most northern Great Plains states and provinces because of perceived rarity and/or insufficient data on its popula- tion status and breeding biology (Anonymous 1991; Rosenfield et al. 1991). Increases in woodland habitat across the region during the mid- to late 1900s have influenced the abundance and distribution of several raptor species (Houston and Bechard 1983; Murphy 1993; Sargeant et al. 1993), and may have also influ- enced the Cooper’s Hawk, e.g., breeding by the hawk now is confirmed or suspected in much of southern Saskatchewan (Smith 1996:103). In eastern North America, the Cooper’s Hawk is classified as an area- sensitive, forest-interior species (reviewed in Grant and Berkey 1999). Recent data suggest a more flexi- ble nesting habit, possibly enabling the hawk to nest successfully in what might appear to be marginally suitable locales (e.g., Rosenfield et al. 1995; Boal and Mannan 1998). To address geographic gaps in knowledge of the species’ breeding status, we mea- sured breeding density and reproductive success of Cooper’s Hawks in several woodland types charac- teristic of the northern Great Plains. Study Area and Methods We studied Cooper’s Hawks across the Souris River basin (48°40’ N, 101°25’ W) of north-central North Dakota. The Souris River forms a 110-km loop extending from Canada into North Dakota (Figure 1). During 1994-1995 and 1998-2000, we located nests of Cooper’s Hawks along the western half of this riverine loop, including the Des Lacs River and inter- mittent tributaries (“coulees”) of both rivers (Figure 1). This area included parts of Des Lacs National Wildlife Refuge (NWR) and Upper Souris NWR in Ward, Burke, and Renville counties, and privately- owned woodlands in Ward and southwestern McHenry counties. Woodlands occurred mainly within river floodplains and on north- and east-facing slopes (5—25°) of coulees, and were dominated by Green Ash (Fraxinus pennsylvanica) and American Elm (Ulmus americana). We defined woodlands within floodplains as floodplain woodlands, and those in adjacent coulees as coulee woodlands. Both woodland types were narrow (< 0.2 km wide) and lin- ear. The area within | km of the Des Lacs and Souris rivers was composed of approximately 14% wood- land, 35% grassland (native prairie and introduced grasses), and 31% cropland. Palustrine wetlands, 580 2002 Hl Intensively searched areas [__] National Wildilife Refuge (NWR) NENNEMAN, MuRPHY, AND GRANT: COOPER’S HAWKS IN THE GREAT PLAINS 581 40 Kilometers FIGURE 1. Cooper’s Hawk study area in the Souris River basin of north-central North Dakota during 1994-1996 and 1998-2000, and two intensive study areas: A = 1998 intensive study area (3951 ha), containing 15 nest areas (10 occupied in 1998); and B = 1996 intensive study area (5263 ha), containing 18 nest areas. open water, and rural towns and cities each composed < 10% of this riverine landscape. During 1996 and 1998-2000, we located nests of Cooper’s Hawks along the eastern half of the Souris River loop in North Dakota, on the southeastern third of J. Clark Salyer NWR in McHenry County (Figure 1). Floodplain woodland along the Souris River at J. Clark Salyer NWR was dominated by Green Ash and American Elm and was interspersed with sedge (Carex spp.)/grass (Gramineae) mead- ows. Uplands at J. Clark Salyer NWR were sandy, rolling hills with plant communities composed of 58% native prairie and 42% broadly scattered wood- lands dominated by Quaking Aspen (Populus tremu- loides), Bur Oak (Quercus macrocarpa), and Balsam Poplar (P. balsamifera). We defined these as sand- hill woodlands. Climate of the study area was semi- arid to subhumid continental with annual precipita- tion averaging about 42 cm (U.S. Fish and Wildlife Service, unpublished data). Reproductive success We located Cooper’s Hawk nests during late April through June by searching woodlands throughout the study area on foot, including 14 nest areas recorded during 1986-1987 (G. Berkey and R. Martin, unpublished breeding bird atlas for Ward County; U.S. Fish and Wildlife Service, unpublished data). Because woodlands on our study area were either linear (coulee and floodplain woodlands) or in discrete groves (sandhill wood- lands), we could scan all trees for stick nests by slowly walking transects through each woodland. As an aid in detecting nesting hawks, we broadcast a Cooper’s Hawk alarm call every 10-20 min from a hand-held cassette player (Rosenfield et. al. 1985). A nest was considered occupied if eggs were laid, and successful if it produced at least one large nestling (2 23 d old [Steenhof 1987]). We defined a nest area as the area within 0.4 km of the original nest found. A nest area was considered reoccupied if a new nest was found within 0.4 km of where nesting had occurred at least six years previous (Rosenfield et al. 1995). Nesting density We measured Cooper’s Hawk nesting density by selecting two intensive search areas where we had no prior knowledge of the hawk’s breeding abundance other than observations of two nest areas on each search area. The two intensive search areas were selected because they afforded ready legal access and an arbitrary boundary (e.g., refuge fences) to confine our search. Also, a basic inventory of breed- ing raptors was among the top priority information needs on area NWRs. Although woodland types on the two intensive search areas were representative of those available on the study area, we acknowledge that nesting densities may not have been uniform across the study area. In 1996, we searched a 5263- ha area in the sandhill woodlands at J. Clark Salyer NWR (Figure 1). We did not include floodplain woodlands in our intensive search area at J. Clark Salyer NWR because flooding precluded access to most of this area through late spring, and we were not confident all nests in the multi-storied canopy of floodplain woodlands would be detected after leaf- out. In 1998, we searched a 3951-ha area on Des Lacs NWR (Figure |). We measured the distance between Cooper’s Hawk nests on aerial photographs 582 (1:7920, 1:15840) to determine nearest-neighbor nest distance. While on intensive search areas during spring and summer, we also recorded locations of occupied nests of other diurnal raptor species to assess abundance of Cooper’s Hawks relative to other breeding raptors. Results Reproductive success Cooper’s Hawks returned to the study area in mid-April, and most females initiated egg-laying in mid-May (mean initiation date based on backdating from estimated nestling ages = 16 May, SE = 0.8 d, n= 57). We identified 56 Cooper’s Hawk nest areas in the study area during 1994-1996 and 1998-2000. Eleven of 14 (79%) nest areas occu- pied in 1986-1987 were reoccupied when first checked in 1994-1995, and 10 of 12 (83%) nest areas occupied in 1994 were reoccupied in 2000. Reproductive success was recorded for one year only at each of 31 nest areas, for two years at 15 nest areas, for three years at six nest areas, and for four years at four nest areas. Cooper’s Hawks pro- duced an average of 2.0 large young/occupied nest and more than two-thirds of nests were successful (Table 1). However, Cooper’s Hawks were less successful when nesting in coulee woodlands com- pared to sandhill woodlands (x? = 6.783, df = 2, P = 0.034; Figure 2). Including one pair that success- fully renested after a failed nesting attempt, 15 of 30 (50%) nest failures occurred during the egg stage (mid-May through mid-June), five (17%) dur- ing the nestling stage, and timing of 10 failures (33%) was unknown. Shell fragments found at seven nests depredated during incubation were con- sistent with shell fragments found at duck nests depredated by Raccoons (Procyon lotor; Sargeant et al. 1998). Two failed nests contained eggs with small (1—1.5 cm) punctures attributed to avian predators, but the eggs may have been destroyed after nest abandonment. One nest failure was attributed to nestling predation by a Great Horned Owl (Bubo virginianus), and a second nest failed THE CANADIAN FIELD-NATURALIST , Vol. 116 - during incubation when the female was killed on the nest, probably by an avian predator. Nesting density Ten nests were found on the intensive search area at Des Lacs NWR in 1998, yielding a nesting density of one occupied nest/395 ha. Eighteen nests were found on the intensive search area at J. Clark Salyer NWR in 1996, yielding a nesting density of one occupied nest/292 ha. Nearest neighbor distance between nests averaged 2.1 km on Des Lacs NWR (range 1.3—3.2 km) and 1.5 km on J. Clark Salyer NWR (range 1.0-2.3 km). Cooper’s Hawks were the most abundant nesting falconiform we observed on intensive search areas (Table 2). Northern Harriers (Circus cyaneus) may have been nearly as common at Des Lacs NWR (at least one occupied nest/ 439 ha), but we were unable to locate all Harrier nests because our survey methodology was better suited for tree-nesting raptors. We were confident that the Harrier was an uncommon nesting raptor (only one or two nesting pairs) in the sandhill wood- lands at J. Clark Salyer NWR. Discussion Reproductive success Reproductive success in our study (2.0 young/ occupied nest and 69% nest success) was in the mid- dle of ranges reported for other regions (1.6—2.8 and 53-85%), as summarized by Rosenfield and Bielefeldt (1993). Our reproductive success data, like those from a prairie-parkland area in northwest- ern North Dakota (2.4 young/occupied nest [Murphy 1993]), suggest that Cooper’s Hawk reproductive success in North Dakota is comparable to that of conspecifics in other regions. Additionally, observed nest area reoccupancy of 79% and 83% across six to nine years suggests a relatively stable population (Rosenfield et al. 1995). Differences in woodland configuration may explain differences in nest success among the three woodland types we studied. Cooper’s Hawk nest success in coulee woodlands may have been lower TABLE 1. Cooper’s Hawk reproductive success in north-central North Dakota by woodland type and for all nests com- bined, 1994—1996 and 1998-2000. Numbers presented are mean + SE (number of nests). Woodland type Reproductive measure Coulee Floodplain@ Sandhill All nests® Mean clutch size 3:8 = 0:2 (5) 2.0 + 0.0 (1) 3.4 + 0.2 (18) 3.5 + 0.2 (28) Mean number of large young/occupied nest* 1.6 + 0.2 (44) 2.4 + 0.4 (18) 2.4 + 0.2 (28) 2.0 + 0.2 (96) Mean number of large young/successful nest¢ 2.8 + 0.2 (25) 3.30.2 (13) 2.8 + 0.2 (24) 3.0 + 0.1 (66) % Nest success 57 + 7.6 (44) 72 + 10.9 (18) 86 + 6.7 (28) 69 + 4.8 (96) Includes one failed nesting attempt that was followed by successful renesting. ‘Includes one nest in a tree planting (4 year), and two nests in urban parks (1 year each). ‘Includes two nests classified as successful when young were 16- to 18-days old, because subsequent nest visits were not made. 2002 sandhill n=28 floodplain coulee nieAe n= 44 Percent nest success (95% confidence interval) coulee floodplain sandhill Woodland type FIGURE 2. Cooper’s Hawk reproductive success in three woodland habitat types in north-central North Dakota, 1994-1996 and 1998-2000. than in other woodland types because the narrow coulee woodlands act as travel corridors for preda- tors. Vander Haegen and DeGraaf (1996) found that artificial nests in narrow riparian buffer strips (20-40 m and 60-80 m wide) were more likely to be depredated than those in more extensive riparian forests (continuous forest extending > 400 m from the river). In our study, Cooper’s Hawks nested most successfully in sandhill woodlands where predators (e.g., Raccoons) may have been less common and less able to locate nests among the widely scattered patches of woodland. Floodplain woodlands, although somewhat narrow like coulee woodlands, were surrounded by about twice as much woodland cover as were coulee woodlands (Nenneman et al. unpublished data), which may explain the intermedi- ate nest success we observed in the floodplain. Nesting density Nesting densities in this study (one occupied nest/292-395 ha) match the highest densities reported for the species (one occupied nest/ 272— 437 ha), in rural and urban Wisconsin (Rosenfield et al. 1991, 1995) and urban Arizona (Boal and Mannan 1998). Nesting densities in other parts of the northern Great Plains have been much lower: one occupied nest/1918 ha for mixed deciduous- NENNEMAN, MURPHY, AND GRANT: COOPER’S HAWKS IN THE GREAT PLAINS 583 coniferous woodlands of the Little Missouri River badlands in southwestern North Dakota (point esti- mate, 1978 stratified random sampling [Postovit 1979]), and one occupied nest/2174 ha for a prairie-parkland refuge in northwestern North Dakota, where woodland cover comprised only 2% of the area (Murphy 1993). Reynolds and Wight (1978) reported a nesting density of one occupied nest/1857 ha in contiguous coniferous forest in Oregon. Results of our study (sandhill woodlands at J. Clark Salyer NWR) and Rosenfield et al. (1995) suggest that Cooper’s Hawks may reach highest nesting densities where woodlands make up roughly 40% of the landscape. Nesting densities, reproductive success, and nest area reoccupancy observed in our study suggest Cooper’s Hawks can maintain viable breeding popu- lations in at least some contemporary woodlands of the northern Great Plains. Cooper’s Hawks likely are increasing in the region because woodland cover has expanded since the early 1900s due to fire suppres- sion, extirpation of bison, and planting of tree shel- terbelts (Sargeant et al. 1993). For example, both intensive search areas in our study were nearly devoid of trees and presumably nesting Cooper’s Hawks only 90-130 years ago (U. S. Fish and Wild- life Service, unpublished data). Relative to other regions, Cooper’s Hawk nesting habitats in North Dakota differ structurally and are fragmented (Nenneman et al., unpublished data). Cooper’s Hawks also are nesting in other newly available habitats in the state, such as tree shelterbelts at iso- lated farmsteads (R. Murphy, unpublished data). Houston and Bechard (1983) indicated that Red- tailed Hawks were limited in the prairie of southern Saskatchewan by lack of trees for nesting, but have become abundant since settlement in the late 1800s as woodland cover has increased. Our findings sug- gest that Cooper’s Hawks in North Dakota had been similarly limited by the lack of trees for nesting, rather than by food resources, and have increased in abundance with increased woodland cover. Cooper’s Hawks are flexible in their selection of nesting habi- tat, and readily nest in woodlands available in North Dakota. Resource personnel should be aware that this species likely is a significant component of TABLE 2. Nesting densities of Cooper’s Hawks relative to other diurnal raptor species on 5263- and 3951-ha intensive study areas at J. Clark Salyer NWR and Des Lacs NWR in North Dakota, 1996 and 1998. Di a Species Cooper’s Hawk, Accipiter cooperii Red-tailed Hawk, Buteo jamaicensis Swainson’s Hawk, B. swainsoni J. Clark Salyer NWR Density as ha/occupied nest (71) Des Lacs NWR 395 (10) 659 (6) 659 (6) 292 (18) 478 (11) ae (QO) ——EE —__——_________.e aNone observed. 584 breeding raptor communities in areas of the northern Great Plains where deciduous woodlands are an important (> 10%) landscape component. Acknowledgments G. Berkey and R. Martin kindly provided Cooper’s Hawk nest area locations for Ward County from 1986-87. We thank G. Berkey, E. Madden, A. Eddingsaas, D. Guenther, G. Wolf, and especially R. Martin and M. Sondreal for help with nest searches. E. Madden, E. Merrill, and M. Restani commented on early manuscript drafts. Reviews by C. Boal, S. DeStefano, A. Erskine, R. Mannan, and J. Squires helped improve the final draft. R. Rosenfield provid- ed valuable advice throughout the project. During the 1995 field season, MPN was an undergraduate intern under auspices of the College of Natural Resources, University of Wisconsin-Stevens Point; support was provided by Des Lacs NWR Complex and the Challenge Cost Share Program of the U.S. Fish and Wildlife Service (FWS). FWS (Des Lacs NWR Complex, J. Clark Salyer NWR Complex, Upper Souris NWR), North Dakota Game and Fish Department’s nongame program, and North Dakota Falconers’ Association cooperatively funded this research in 1996. FWS helped support the work in 1998-2000. Literature Cited Anonymous. 1991. Status of endangered wildlife in Can- ada. Committee on the Status of Endangered Wildlife in Canada (COSEWIC), Ottawa, Ontario, Canada. Boal, C. W., and R. W. Mannan. 1998. Nest-site selec- tion by Cooper’s hawks in an urban environment. Journal of Wildlife Management 62: 864-871. Grant, T. A., and G. B. Berkey. 1999. Forest area and avian diversity in fragmented aspen woodland of North Dakota. Wildlife Society Bulletin 27: 904-914. Houston, C.S., and M. J. Bechard. 1983. Trees and the Red-tailed Hawk in southern Saskatchewan. Blue Jay 41: 99-109. Knopf, F. L. 1996. Prairie legacies — birds. Pages 135-148 in Prairie conservation: Preserving North Amer- ica’s most endangered ecosystem. Edited by F. B. Sampson and F. L. Knopf. Island Press, Covelo, Cali- fornia. Murphy, R. K. 1993. History, nesting biology, and preda- tion ecology of raptors in the Missouri Coteau of north- western North Dakota. Ph.D. dissertation, Montana State University, Bozeman, Montana. 212 pages. Postovit, H. R. 1979. Population estimates of breeding raptors in the North Dakota Badlands. M.S. thesis, North Dakota State University, Fargo, North Dakota. 50 pages. THE CANADIAN FIELD-NATURALIST EO i SS OE DON Vol. 116 - Reynolds, R. T., and H. M. Wight. 1978. Distribution, density, and productivity of Accipiter hawks breeding in Oregon. Wilson Bulletin 90: 182-196. Rosenfield, R. N., and J. Bielefeldt. 1993. Cooper’s Hawk (Accipiter cooperii). In The Birds of North America, number 75. Edited by A. Poole and F. Gill. The Birds of North America, Inc., Philadelphia, Pennsylvania. 24 pages. Rosenfield, R. N., J. Bielefeldt, J. L. Affeldt, and D. J. Beckmann. 1995. Nesting density, nest area reoccu- pancy, and monitoring implications for Cooper’s Hawks in Wisconsin. Journal of Raptor Research 29: 1-4. Rosenfield, R. N., J. Bielefeldt, R. K. Anderson, and J. M. Papp. 1991. Status reports: Accipiters. Pages 42-49 in Proceedings of the midwest raptor management symposium and workshop. Edited by B. E. Pendleton and D. L. Krake. National Wildlife Federation Scientific and Technical Series 15, Washington, D.C. Rosenfield, R. N., J. Bielefeldt, R. K. Anderson, and W. A. Smith. 1985. Taped calls as an aid in locating Cooper’s hawk nests. Wildlife Society Bulletin 13: 62-63. Sargeant, A. B., R. J. Greenwood, M. A. Sovada, and T. L. Shaffer. 1993. Distribution and abundance of predators in the Prairie Pothole Region that affect duck production. U.S. Fish and Wildlife Service Resource Publication 194. 96 pages. Sargeant, A. B., M. A. Sovada, and R. J. Greenwood. 1998. Interpreting evidence of depredation of duck nests in the Prairie Pothole region. U. S. Geological Survey, Northern Prairie Wildlife Research Center, Jamestown, North Dakota, and Ducks Unlimited, Inc. Memphis, Tennessee. 72 pages. Semenchuk, G. P. 1992. The atlas of breeding birds of Alberta. Federation of Alberta Naturalists, Edmonton, Alberta. 390 pages. Smith, A. R. 1996. Atlas of Saskatchewan birds. Sas- katchewan Natural History Society, Regina, Saskatche- wan. 456 pages. South Dakota Ornithologists Union. 1991. The birds of South Dakota. Northern State University Press, Aber- deen, South Dakota. 411 pages. Steenhof, K. 1987. Assessing raptor reproductive success and productivity. Pages 157-170, in Raptor management techniques manual. Edited by B. A. Giron Pendleton, B. A. Millsap, K. W. Cline, and D. M. Bird. National Wildlife Federation Scientific and Technical Series 10, Washington, D.C. Stewart, R. E. 1975. Breeding birds of North Dakota. Tri- College Center for Environmental Studies, Fargo, North Dakota. 295 pages. Vander Haegen, W. M., and R. M. DeGraaf. 1996. Predation on artificial nests in forested riparian buffer strips. Journal of Wildlife Management 60: 542-550. Received 9 April 2001 Accepted 20 September 2002 The Forests of Prince Edward Island: A Classification and Ordination Using Multivariate Methods D. G. SoBEY! AND W. M. GLEN2 School of Applied Biological and Chemical Sciences, University of Ulster, Jordanstown, Northern Ireland BT37 OQB. (Research Associate of the Institute of Island Studies, University of Prince Edward Island, Charlottetown, Prince Edward Island) 2W. M. Glen, Natural Resources Division, Department of Agriculture and Forestry, PO Box 2000, Charlottetown, Prince Edward Island C1A 7N8 Canada Sobey, D. G., and W. M. Glen. 2002. The forests of Prince Edward Island: A classification and ordination using multi- variate methods. Canadian Field-Naturalist 116(4): 585-602. In 1991 data were collected on the ground flora in 1200 4 m? plots at 240 randomly selected forest sites on Prince Edward Island. The nearby trees and shrubs, and attributes of the surrounding forest stand were also recorded and soil samples were collected for analysis. Additional topographic, historical and soil data for each sampling point were obtained from maps and a 1935 aerial photographic survey. 1127 of the plots were subjected to multivariate analysis (a TWINSPAN classifica- tion and a DECORANA ordination). On the basis of their ground flora composition, TWINSPAN divided the plots into eleven ground flora community-types, which after further comparative analysis of their tree canopies, stand properties and environmental factors, were assigned to five forest-types, each characterised by particular tree species and soil drainage properties: (1) a wet species-rich woodland, (2) upland hardwood forest, (3) Black Spruce forest, (4) old field White Spruce woods, and (5) disturbed, mainly conifer-dominated, forest. The first three were considered to be heavily modified descendants of pre-settlement forest-types, while the two latter appeared to be largely the products of successional process- es resulting from human disturbance and forest clearance. The ordination indicated that the most important factors respon- sible for the ground flora communities and forest-types on Prince Edward Island are soil drainage and human-associated disturbance. Key Words: forest ground flora, forest classification, forest ordination, Prince Edward Island, TWINSPAN, DECORANA. Prince Edward Island, comprising 5750 km7, is the most agricultural and least forested of Canada’s provinces. Lying within the so-called boreal-broadleaf ecotone (Scott 1995), it contains both deciduous hard- wood forest, and tree species and communities char- acteristic of the more northerly boreal coniferous for- est (Erskine 1960). Since the beginning of European settlement in 1720, and especially in the nineteenth century, its forests have been subjected to large-scale clearance such that by 1935 only 32% of its area was still under forest (Glen 1997), though by 1990 this area had increased to 49% (Anonymous 1992). However, the composition of its forests, either past or present, has never been studied in detail, with no com- prehensive field-based study ever having been carried out. Those studies that have touched on the island, have either been general and brief, serving primarily to place its forests in the context of continental or regional forest-types (Halliday 1937; Rowe 1959; Loucks 1961), or, with forest description being peripheral to other aims, have been entirely qualitative and descriptive (Stilgenbauer 1929; Erskine 1960; MacDougall et al. 1988). The study reported here rectifies this deficiency. It stems from an opportunity that arose in 1991 when the P.E.I. Forestry Division was planning the field work connected with a comprehensive inventory of the island’s timber stock. Under the Division's con- servation remit, it was decided to also collect informa- tion on the ground vegetation at each of the invento- ry’s 1200 sampling points. Because of limitations in time and resources the ground flora assessment at each sampling point was confined to a single 4 m? cir- cular plot, which had the potential for limiting the use- fulness of the data collected. However, as will become evident from the results, even small plots, provided there are many of them, can yield valuable results. Multivariate analytical methods were used to anal- yse the data, the aim being to classify and ordinate (i.e., arrange along gradients of changing plant com- munity composition) the 1200 plots on the basis of their ground flora composition. The plot groups and gradients emerging could then be assessed in terms of environmental and other data recorded for each of the sampling points (either in the field or from maps); e.g., tree canopy properties, topographic attributes, soil properties, and management and historical fac- tors. It was hoped that the analysis would lead to a precise documentation of the variation in the forests of the island and a greater understanding of the environmental factors determining the variation. This study is unusual in that it aims to provide a comprehensive and precise description of the forests of an entire geographical entity. Also unusual in Canadian terms, is the fact that virtually all of this for- est has been heavily affected by 150-270 years of human interference and disturbance. However, this does not mean that the study has no relevance outside wn 586 of Prince Edward Island: the island contains forests belonging to two continental forest types (the northern hardwoods and the boreal coniferous forest) and thus presents an opportunity for a detailed spatial analysis of these forest types in an area where they overlap. Also, at a regional level, the island’s forests show affinities with the forests of adjacent parts of western Nova Scotia and southern and eastern New Brunswick (e.g., see Rowe 1959; Loucks 1961). Methods Field methods The data on which this paper is based were collected as part of the 1990-1992 Prince Edward Island Forest Biomass Inventory, the field work for which was carried out by two teams of workers between June and September 1991 at 240 forest sites (Figure 1) selected prior to the field visits from maps using a fully randomised procedure. At each site, five sampling points were positioned (usually at 100 m intervals) along a cruise line whose compass-bearing and starting point had also been determined prior to the field visit and marked on aerial photographs (1:17,500) to aid location. To enhance inclusion of different forest types the cruise lines were chosen to run across map contour lines, where such occurred. The first sampling point was usually 100 m in from e . \ WESTERN THE CANADIAN FIELD-NATURALIST a a Vol. 116 - the wood edge. Alder scrub and clear-cut areas were to be excluded in the actual sampling and no sampling point was to be closer than 15 m to a boundary between two distinct forest types. With five sampling points at each of the 240 randomly chosen sites, a total of 1200 sampling points were assessed in the field inventory. Site and forest attributes — At each of the 1200 sampling points the assessors recorded the slope (as a percentage) and the aspect (in eight directional classes), and the following standard forest inventory parameters based on a subjective impression of the general area around the sampling point (see Haddon (1989) for a definition of the terms): (1) the forest type; 1.e., whether predominantly hardwood (HH — >75% hardwood, HS — 50-75% hardwood) or soft- wood (SS — >75% softwood, SH — 50-75% soft- wood); (2) the broad age class of the trees (regenera- tion, immature, mature, overmature, uneven-aged); (3) the stocking class of the trees (as four quartile percentage classes); (4) where evident, the origin and management history of the stand (i.e., swampy, bog, burn, partial cut, thinning, clear-cut, plantation, old field, or a combination of these); and (5) evidence of damage due to logging, insect defoliation, disease and windfall. EASTERN FiGURE |. The 240 randomly-selected sampling sites assessed in the 1991 field survey — there were five 4 m* ground flora plots at each site. The three regions into which the island was divided for the purpose of this study are shown. 2002 1127 595 165 430 6 159 79 351 oe | [| | 2 Gp. Gp a 4 Wet Upland Rich Hardwood Woodland Forest SOBEY AND GLEN: FORESTS OF PRINCE EDWARD ISLAND 587 532 439 93 283 156 41 52 FiGURE 2. The TWINSPAN classification of 1127 ground flora plots: dendrogram showing the dichotomies leading to the fifteen plot groups generated at the fourth division level, and the five forest-types to which the plot groups were assigned. (The number of plots at each division is shown.) Soil properties — A soil pit was dug near the sam- pling point and using standard soil survey techniques the following physical properties were determined in the field: soil drainage class (well-drained, moderately well-drained, imperfectly-drained, poorly-drained); rooting depth (cm); thickness of the A horizon (cm) (i.e. the distance from the start of the mineral soil under the H horizon, to the B horizon); root restriction factors (at all sampling points recorded as either due to a high water table or to soil compaction); and the tex- ture of the B horizon. Soil samples were taken from both the H horizon (from just above the mineral soil) and from the B horizon (so as to include both the upper and lower part of the B). These were subsequently analysed for pH, total carbon and nitro- gen, and available levels of phosphorus, potassium, calcium, magnesium and sulphur, as well as micronutrients. The chemical analyses were carried out by the Prince Edward Island Department of Agriculture and Forestry Soil and Feed Testing Laboratory in Charlottetown using the following methods: the soil samples were oven-dried overnight at 40°C, ground in a soil crusher and passed through a 2 mm sieve. The pH was determined electrometrically on a 1:1 volume ratio of the ground and sieved soil and distilled water; total carbon and nitrogen were determined by com- bustion on a CHN analyser; available Ca, Mg, K, P and S were extracted using a Mehlich III extract at a 1:10 soil to extract ratio, shaken and filtered, and run on an Inductively Coupled Argon Plasma (ICAP) Spectrometer. For carbon and nitrogen, final values were expressed as percentage of oven-dried soil, for all of the other elements, as parts per million. Vegetation assessment — At each sampling point the vegetation was assessed as three separate cate- gories: 1. Trees (=> 9 cm DBH, including standing dead trees) were assessed using a ‘point sampling’ tech- nique (Watts 1983), which involved the use of a ‘vari- able-radius plot’ centred on the sampling point, with trees being selected for inclusion in the sample using a wedge prism of Basal Area Factor = 2 m’. The species of each tree and its DBH to the nearest cm was recorded. From this data, the percentage contribution (to the nearest 10%) of each tree species to the total woody biomass in each variable-radius plot was cal- culated. Two trees, generally the largest broadleaf and conifer, were selected for measurement of height (using a clinometer), and age at breast height (using an increment borer). 2. Small trees and woody shrubs (2 1.3 m in height but <9 cm DBH) were assessed within a 16 m? circu- lar plot (radius 2.26 m) centred on the sampling point. For each plant, the species and vitality (i.e. whether living or dead) was recorded, as well as its DBH to the nearest cm. 3. Ground flora — all species of vascular ground plants including woody shrubs < 1.3 m in height (but excluding tree species); and selected species of moss- es, liverworts and lichens were assessed within a cir- cular 4 m? plot (radius 1.13 m), the centre of which, to avoid the effects of the trampling involved in the tree and shrub sampling, was moved 5 m onward from the sampling point along the bearing of the cruise line. Based on a visual impression of its cover-abundance, each species or other taxonomic category was given a percentage cover value to the nearest 10% with a minimum of 5%. 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(¢ dnoip dS.) }SO10,J SpOOM SO10J }SO10F pur]poon sonidg sonids JoyIuod poompieH yory yor uM poginisiq pueydy JOM (sdnois NVdSNIML Sutpuodsaiios pur) sadA [.-LSAYO] ["9}0U]00} das — js9} UOSLIedWOD a[dn[NW gJJoYdS dy) 0} JOJO ONTVA YORI BUIMOT[OJ SONI] YL] ‘(e) JOP & AQ PayeoIpUT SI SseUIOIG %]| > 1v BdUasaId saisadg = sadA}-Jsa10} NVdASNIAAL 94} JO sod snipes-s]qeieA oy) UI sseUIOIG 992) [k10} BY) 0} saIdads dan jedrouLd ay) Jo UONNGLNUOD aSeIUAdI0d oy) Jo (GS F) URIWI OU], “| ITAVI 589 FORESTS OF PRINCE EDWARD ISLAND SOBEY AND GLEN 2002 ‘(S9LI9s [10S JOJ p pur ‘sid [10s 10} ¢ = “J"p ‘SUNOS Joy ‘by = "JP ‘SMOI JOJ) 10'0 > d ye jueoriusis ore pjoq ur payutd suns -X Mos JO ULINTOD ‘ssejo oseureIp pure odA}-jsoI0] se[NoNed oy UZBMIJ9q UONPIDOSSE dy) JO sINIeU JY) Sa}edIpUT .-, JO .+, @ sadejusosad yons 10,4 (1000 > d - xxx S10'0 > 4 - xx 'S0'0 > d - x ‘LT = JP) prog ur poyunid ore JuRoTpIUsIS AT[LONSNeS d19M sonjea -X Surpuodsauos ay yotym Joy sade ‘uaoied BYL “(L670 = A S.touresD) :(100'0 > d ‘91 = FP) L'E8e = 2X [P10 :saqaas [los “(LT p'0 = A S.soweID) (100'0 > d ‘ZI = FP) O'S9S = -X [e101 ‘syd [IOs :uoNPIDOsSsE Jo s}sa -X4 oo eee ee OT i OOO —_00 OO C801 08 Cel £87 8CV 9S I SLNIOd dO YasWON a pe ee ee SS ee eee 0°L9 P'S¢e rol €ril S'9ST SUNS ULINJOS -X nO eA en as a Se ee ee eee 9°98T CEC xk + OSS *«k— 9°6 L'VC xk — O'L xxx + £°09 pouresp-AjJ00g Cee Vol x + 88ST x— UP DET * — $°S xxx + 6°61 poureip-Apoopoduy 9 Le ag O'€ ae VV 790 pouresp-|[am Ajayesopoyy ULI C8 x — 8°8 ees x — 96€ xxx + OTL xxx — Ll POUTeIP-[[O A Orv 6 v1 OT xk + 9°67 * + S'6L «—L'0r xx — SP poureip-A[pidey ‘SdVJ SAIMAS TOS (q) eee 5 sn a ee ee eee eee Tvel 6'0€ Ltr €'O7I L’S€Z suns UUINIOS -X Pe SE EE a ae eae ene ee NE Peo eM eT SME EL sss T9OT¢e COL xxx + OOP «xk — 0 ek — US xx — €7°O xx + OCP poureip-Aj100g 0°6L 9°8 aa + €°97 ae — ST ax + SET xx — OT «x + O°9T pouresp-Apoajioduy] 69¢ een 881 [vl xz + OLT kk — 9°6 Lt poureip-|[am Ajayeapoyy] Leer 89 +x — OST xk + PDS c9S xxx + 9°88 xx — 6OLT POUIeIP-[]O MA ‘SLId TOS (B®) eee ——————————————— oe se SS SS ee suns sJUIOg (pI + €I (OL (6+8 (L+9+6 SSVIZ-D ADVNIVUG TOS MOL IV sdnoip dL) dnoip dS.) sdnoip dS.L) sdnoip dS§L) (¢€ dnoiy dS) 2X }SOI0J SPpOOMA }S910,] S104 pur[poom sonids sonids JoyIuog poompiey yory yr ov. poqinistq purjdy JOM (sdnois NVdSNIML Sulpuodsazioo pur) sadA J -LSadO4 —oows———————ooOoOoOoOoOoOoOoOO019namomaRDD9O99S9MOOwOwOwqwomomwmronmwmaqyqywq*nang0»s«=>$="—_ OO EEEE—S=~<~ ey Wa ie ns ets tidal rivers and estuaries. a 2 aS RofotSaSlon |" | a 2. Distance from nearest road or building — the 77) er —s tee E é ‘ A Sa] 8 A = ae aoe a a t vA 2 minimum straight-line distance (m) from the sampling Ge oO 5 og 41 : Salil s = 5 point to the nearest building or road (whichever was > cle = nearer), not counting as roads: “trails, cut-lines or 3 v 99 ee x 5 > portages”. _ 5 = - P a Bi he ae ‘o Ss . Elevation above sea level — as estimated from E All = Seems oe SS i a 3. Elevat. b level timated fi 2Ele|2s z Sp SSSE = = ol = map contour lines (25 foot intervals), the values being 25] 2 ae soe SSL SHZS HIG] & converted to metres. —_ _ ~~ — fon) ~— ‘2 — a A &E 5 Y= Sree eye = = x » 4. Size of wooded area — the area (km?) of the 43 S ST SRE SH woodlot or forest as determined from the 1:50,000 50 = maps, on which wooded areas are indicated by green a == ! EL 3 shading. 5 a Pee S An additional parameter was obtained from a map 2) 5 ; uo) : ar 2 2 9 % i} in MacDougall et al. (1988): =* 2. — S OE ae CS 3S = ; ; ; 52 ge |Saeasoaa 8 5. Corn heat units — an indicator of summer heat 2 et ~ oo) an Ss = 5 5 F zZ¢ = 3 3 5 eas ] aa = Sar Vigie based on cumulative daily temperatures in the grow- u ioe) a So SS C0 . ‘ . = & 74 2 A. widataondg = ing season. The corn heat units for each sampling ee Ww = . . . o 5 =) ae eS point were read from contour lines in Figure 2 of oS S > a MacDougall et al. (1988). x vo o. . i= . In addition, two parameters were obtained by com- me = puter from the Geographic Information System 4 2 3 (G.I.S.) database for the province: a5 ee tes 6. Status in 1935 — the status of the sampling = ¢ FE & |§ 6 points in 1935 at the time of the first aerial photo- 2s a > SES elall = graphic survey of the island. The forest boundaries in e 2 si SS < ¢ 3a5 & S|] % the aerial photographs had been transferred by peas’ a coe a “ 6 5 cs a Forestry Division photo-interpreters to maps which ae i ~ £4 soesl2i z were subsequently digitised and incorporated into the KS FIROZOROSGIZIE G.I.S. system (Glen 1997). The database was queried NO i=) S iw) SOBEY AND GLEN: FORESTS OF PRINCE EDWARD ISLAND 591 ey pleonmaenm fe 2. for the status of each of the 1200 sampling points in a BE SOR ASA z g 22 ~ 1935, with their status being recorded as either forest- 8 - Bick, 2 a5 covered or cleared land. = re RON is a ien ree 2 7. Soil series — as defined by the Prince Edward 2 aaeleransnen |Siis ge Z Island Soil Survey (MacDougall et al. 1988). This was = §& a. 53 obtained from 1:10,000 soil series maps produced by a Le 2% %%% __ the Soil Survey, as digitised and incorporated into the e oo BS 63% GS. database. Where a map unit was listed as com- § £ on % # Se &2o prising more than one soil series, only the first named z 82 Eley age Eo a 5 = @ series was recorded. ee a ae =e a S = S re = . 3 S & Bg Methods of data analysis ‘3 = a of aed Two multivariate methods were used: Two-way Ss me S5 2 2 = Indicator Species Analysis (TWINSPAN or TSP), a 6 ie 22 5&4 classificatory technique, and Detrended Correspon- = 4 ao 38 & & dence Analysis (DECORANA or DCA), an ordina- 8. Se |, ig <= & 8% _ tion technique. Both have been specifically developed 2 = a/e x es 5 2. : = for the analysis of plant community data (Hill 1979 a, a hve 5 Brie e ee, S a 1 |e 38 225 b; Gauch 1982) and have been used extensively in for- 3 a. Ae |erarneac 5 S s 2 - Te CSE and other vegetation studies, including several 2 co 2 3 e s De 3 Canadian forestry studies (e.g., Jones et al. 1983; Sims = |lzlz a Ay et al. 1989; Bergeron et al. 1992; Bowling and = aa 3 2s 3 5 Zelazny 1992). They are designed to be used in a = Zio ae oo 2 complementary fashion on the same set of data, thus £ = cape a s a8 2 ° enabling both the recognition of plot groups compris- & ||| Bo _ 2 e 2.2 ing distinct plant community types, and the spatial B ilelse + 1S oo ||/= = 25-22 ordering of these groups and their component plots in B eS slaseshslz|8 | $58 28s f thei II similarity to each other S 5/56 nSsadsss-ls es eso terms of their overall similarity - 2 a Bo = Cee ne 5 TWINSPAN and DECORANA were applied to the a Fle Ee 2 5 S 2 ground flora data using Cornell University computer 6.5/4 aun 128% programs (Hill 1979 a, b) modified to enable the anal- & 5 & E3338: ysis of all 1200 plots in a single run. Seventy-one of Sala) Ze ee ss 8 2 — the 1200 plots were omitted beforehand from the anal- £2) =| 5 = 2 a°8 £3 ysis, 18 in which no ground flora species had been as elStle ‘ Fr 82555 recorded and 53 for which no trees were recorded in Api z|s'o|* ee of> Bo SBS the area of the sampling point. Of the 225 taxa record- q s © @ “ eee = 4 ee . FIZss ams z ed in the Inventory, 13 of the 16 non-specific genera = = = 5 AcyTSod—ls : 2 5 were omitted from the analyses (the generic categories 25 Go Ow Sp $ 2 retained were Amelanchier, Sphagnum, and Mnium). S 5 Cia Ak s > «= — All plant species occurring in fewer than five plots ~ I Ss = ES 23 were also Slee which resulted ¥ of further pros Sé 2) C° &* & being reduced to zero species, and these were also E s 5 oe be ef a 3 ~ omitted from the analysis. In the end, 1127 plots and ae E S * # mice: 5 o28 : a3 sane were subjected to TWINSPAN and DECO- ome) i Lt 1 (hill So see ’ 26|| |San/2cSeeel/s/" |lgSis 222 The DECORANA ordination and TWINSPAN ec wis) e's ene 3 3 8 classification are based entirely on the ground flora Bg S % ran E 2§ data. The second stage of the analysis consisted of the Ete * » 88 interpretation of the products of these analyses (i.e. a oo D oe boy 3 the plot groups and two-dimensional graphs). The < 2 2 3 Tv 4 6 # groups were first described in terms of their ground = E 2 bp gv Z 2s Z flora composition and then each was interpreted using ‘S rs 4 wy = a is 3 -"23 2 the independent environmental data (including data on 6 ‘s = = ch ane 2 Me « a3 = the tree canopy). Differences between the various E », Z ee op B 8 me & £ Zi E a e 5 TWINSPAN groups were compared statistically, as gk g¢ ce ole as 3 = 20 while the scores for the plots on the DECORANA < 2 ¥ Rls Peg a 2S ze cs z 2 42 E ‘3 axes were tested for correlation and/or association z “ z : fs ge E as 3 8 Z lla gs 2 F with the independent euyionnecnial ron ae sec eS ADTIOORMAAZO RIA I+ y+ ond stage of the analysis dBASE IV was used to sort WET RICH WOODLAND DISTURBED CONIFER-DOMINATED FOREST BLACK SPRUCE FOREST the plant and environmental data for the various TWINSPAN groups, and the computer package SPSS (Statistical Package for Social Scientists) (Norusis 1990) was used in the statistical analyses. Among the tests applied were one-way analysis of variance (with the Scheffé multiple comparison test), the x? test of association (including Cramer’s V), and Pearson’s correlation coefficient. After the multivariate analyses had been completed, in order to verify in the field the validity of the TWINSPAN groups, and to record photographically the appearance of their ground flora, in the summer of 1994 (using the same cruise line compass bearings and distances as in the 1991 field survey) the sam- pling points of 26 plots selected as representative of the eleven TWINSPAN groups were visited. THE CANADIAN FIELD-NATURALIST ),—— A Ae ES ey CY Vol. 116 UPLAND HARDWOOD FOREST yah WHITE SPRUCE WOODS FIGURE 3. Maps showing the sampling sites (out of 240) at which each forest-type occurred. A dot at a site indicates that at least one of the five 4 m? ground flora plots at that site was classified in a ground flora community-type belonging to that forest-type. The sampling sites where ground flora plots of the forest-type were not present are visible as smaller points. (See Figure 1 for a clearer indication of the location of all of the sampling sites.) Results TWINSPAN classification of the ground flora plots The TWINSPAN classification of the sample plots to the fourth division level generated fifteen ground flora plot groups (Figure 2). Disregarding the four groups with fewer than ten plots leaves eleven groups, ranging in size from 12 to 196 plots (see Appendix | for the ground flora species frequencies for each group). Subsequent analysis, as outlined below, indi- cates that nine of these eleven groups belong to five forest-types. Examination of the tree canopy composition (Table 1) indicates that the first division largely acts to sepa- rate the plots having a large broadleaved component in their canopies (the 595 plots to the left in Figure 2) from those with a high conifer component (the 532 2002 SOBEY AND GLEN: FORESTS OF PRINCE EDWARD ISLAND 593 plots to the right). The broadleaved plots were then split into two groups, the smaller group (Group 3 — 156 plots) being those of a more base-rich and wetter swamp-type woodland (Tables 2 and 3) in which Red Maple (Acer rubrum) predominates, with American Elm (Ulmus americana), White Ash (Fraxinus ameri- cana) and White Cedar (Thuja occidentalis) occurring at low frequencies (Tables 1). The larger group (430 plots) consists of the dryer or ‘upland’ hardwoods, further divided into three sub-groups (Groups 5, 6 and 7) which appeared to differ primarily in relation to the amount of past human disturbance: Group 5 showed the least disturbance: of the three groups, it had the highest proportion of the tolerant hardwood species (31% of its woody biomass comprised Sugar Maple (Acer saccharum), American Beech (Fagus grandifo- lia) and Yellow Birch (Betula alleghaniensis)), as well as the oldest and tallest trees of any group (see Sobey (1995) for further details). The 532 conifer-dominated plots in Figure 2 were divided into a larger group (439 plots) showing asso- ciations with dryer and less acid conifer woods (Tables 2 and 3), and a smaller group (93 plots), asso- ciated with wetter and more acid Black Spruce (Picea mariana)-dominated woodland. The larger group ended up as four groups, two of which (Groups 8 and 9) were quite similar and appear to represent disturbed conifer-dominated forest on dryer soils (these plots are dominated by Balsam Fir (Abies balsamea) with smaller contributions from Red and White spruce (Picea rubens, P. glauca) (see Table 1); the third group (Group 10) consisted largely of the plots of old field White Spruce woods (Tables 1, 4, 5); and the fourth group (Group 11 with only 21 plots) was a species-poor group, which analysis and field observa- tion (Sobey 1995) indicated to be more closely related to the broad-leaved plots than to the conifer plots. The 93 plots of the Black Spruce-dominated forest ended up as three groups, one (Group 12, with only twelve plots) containing an odd mixture of acid and more nutrient-demanding ground flora species. Analysis and field observation suggests it represents a plot-type transitional between Groups 3 and 13 (see Appendix 1). The other two groups comprise the Black Spruce forest proper, sub-divided into a wetter bog-type for- est (Group 13) and a drier lichen-heath-type forest (Group 14) (see Appendix 1). In summary, the eleven ground flora community- types can be taken to represent five forest-types, each showing its own distribution pattern on the island (Figure 3) (see also Appendix 2 for a summary of the factors associated with each forest-type): a wet species-rich swamp-type woodland (Group 3); upland hardwood forest (Groups 5, 6 and 7); disturbed conifer-dominated forest (Groups 8 and 9); old field White Spruce woods (Group 10); and Black Spruce- dominated forest (Groups 13 and 14). These forest- types show striking differences in geographical distribution (Figure 3), topography (Table 6), soil x2 row sums 55.3 150.6 All Points 7301 26.9 1082 Black Spruce Forest (TSP Groups 13 + 14) 67.5 32.5 le? 80 White Spruce Woods (TSP Group 10) 24.4 — *** 1S ree 162.6 135 Disturbed Conifer Forest (TSP Groups 8 + 9) 74.2 25.8 0.17 283 0.308). The percentages for which the corresponding x? values were statistically significant are printed in bold (d.f. = 1, * - p< 0.05; ** - p< 0.01; *** - p< 0.001). For such percentages a ‘+’ or “—’ indicates the nature of the association between the particular forest-type and the 1935 status class. Column or row x” sums printed in bold are significant at p < 0.01 (for rows, d.f. = 4; for columns, 1). 428 Forest 206.0 (d.f. = 4, p < 0.001), (Cramer’s V Upland Hardwood (TSP Groups 86.4 + ** 15.6=""% 38.7 FOREST-TYPES (and corresponding TWINSPAN groups) 5+6+7) 3.2 Rich Woodland (TSP Group 3 79.5 20.5 156 Wet 2 lumn sums test of association: total x 5 Forest-covered Cleared land NUMBER OF POINTS > STATUS in 1935 x- co TABLE 5. The status in 1935 (as either forest-covered or cleared land) of the sampling points of the TWINSPAN forest-types, based on photo-interpretative analysis of a 1935 aerial photographic survey of the island. For each forest-type the percentage of its sampling points in each category is given. 6 Vol. 11 ALIST DIAN FIELD-NATUR A THE CAN 594 5 Se 5 esses) |e = e SSO > : 3 : SI iz 5 g < = 2 @ Z 5b < ; : 2 < pat B Sas lee z + - a 2 SS is a e _ i bie g ot) i a oO ise} aye a. i 5 1S = 2 2 5 1 flora © Be % DCA Axis und flo = ND ane 5 + fa ro es 2¢/@s2 = Z| 5 | scoond axes, Sa BS) PSs ES ws rdin sD of 3 Diy Bega Sy OPS tS RANA o cond axes. for each zh SO+|zhas | ||s Hn in i of plots =| |3 Fe | 5 a8 a ee td ae S A ed aie : oer sven TWINSPAN group n " leven the e Ee sy ey 5 bles 2 and * 2 S20 operties (Ta d 5). Three 0 i Sauces = inage pr bles 4 an upland 2 Ss |o¢ LAS on = chemical a amient ee rich woodland, forest) = oF tees SE5s/2S55 in manag we ruce ified =| Ble B.S a, eee 2 these forest-typ st and the aie me time of 6| b/F A = IS es s hardwood h a pecunay 960) ied =| = e e ; ’ 0 | z E ognisable likely to hav Erskine 1 d forest Sie & ses st-types li t (compare ifer-dominate the x est- n 1fef- ore ola Mees = of for ttleme d con be m =| See @ ean se disturbe ear to ith the rs) — a) o =) Europ (the ds) app’ ted wil mats a CaP 6 —) r two woo socia $ = ms ot 5 S = oO So N ala & nd the Whi essiona p and fores = 6 | 3 Oe Be ae o a 5 ie of human the groun A ordinatio Pe Or geillic: Te 2 effects ‘A ordination of ted by the DC nvalues of N oO es = (e : € F s 5 S mare axes spi two (with ecologically Sel 23 Z the efi e€ an the $| z 22 s ee found that th tively) gav hip(belies ng 21 6 Hae % 2 it was I respec lations s alo S| — ex i= I di0:5 f the re lot group the aa & wn LPS x, co lle 0.65 an icture oO leven p : igure 5 Z| e Ge a ue ae Qi |} Sb eonimenuine f the ele ile in Fig : <| = aot SESSA IS/ SE sie, ee eee igure 4, while the five for S| alo 8 Be nana e The po in Figure 4, d into lari- n Es = = = Qo mon ae se 3 plots. is shown i combine rts andc : e ee oe = D Ne = as a a 5 these aAXeS ? have been Figure +. we groups 44 am 8 2 en grou above. n the p t ric a s eee es discussed hips betwee 3, the we ther 2 > est-types elationshi ion: Group from the o = 2 fies the interr classificati ly separate 5, 6 and 7) ed ape 2 from the 1 spatial y d roups ( ? s dis- 2 os a dent is largely dwood g the les 2 iG a 4% = dland, 1 land har e so than onifer- 5 =e es Sm Oe ae = ris ps; the three ae and 7 mor f disturbed c more = 3 5 a LSSsoGex a Ras roups; t ition, sO lies Bs go lis fh oian 5 8 rlap in Sage the two eine Group 8 Group 9 s = mM So ae $5 E turbed Gro est also ove s 6 and 7, w Group 12, Ss Fe QI iB = ze j dentine ie gP vised Groupe an S . NE PAs the Blnek mare hs both een Groups 5 a whose species dix 1) does (the Black Spru ite Spruce =~ = = < Appe nd 1 the x A = A | é : es 13 Has Groups 13 en Group a7 groups ras < E 22 2/5] 2 13 ( reach ae of the Sari when the gro fo) fee : v8 nea O « a 5 és ‘ef 8 9.5 S|! & % se up) ema ee 5 indicates 2 a|f 832 a | ib g 13, and a s = = | e = 6 ° : | S| 28 Zg\sice re elaaa66 z adie Ee 2002 SOBEY AND GLEN: FORESTS OF PRINCE EDWARD ISLAND 595 beet 3rd Ath 6 White Spruce woods Black Spruce ey Wet rich woodland DCA Axis 2 soyeno Upland hardwood forest 0 1 2 3 ~ 5 6 DCA Axis 1 FIGURE 5. DECORANA ordination (along the first and sec- ond axes) showing the main cluster of plots for the five principal forest-types. (The two axes have been sub-divided into quartiles to facilitate x tests of association for the qualitatively measured attributes of the plots — see Tables 8 and 9.) similar in species composition are combined into five forest-types, four of the groups occupy reasonably separate areas of the ordination while the fifth (Group 10, the White Spruce woods) shows more overlap. DECORANA ordination: environmental factors To determine whether any of the recorded environ- mental factors showed a relationship with the plot ordination, two types of statistical test were carried out: for the variables measured quantitatively Pearson’s correlation coefficients were calculated between each factor and the plot scores along the axes (Table 7), while for the categorical variables, x? tests were used to examine the association between the attribute classes and axis position when the axes were divided into quartiles (Tables 8 and 9). The analyses indicate that two groups of factors show a strong association with the overall ordination of the plots. One group is connected with soil mois- ture levels: attributes indicative of high soil moisture (i.e., poor and imperfect drainage (as determined from the soil pits), poorly and imperfectly-drained soil series (as determined from soil survey maps), a high water table, level ground) all had significant positive associations with the upper ends of each of the two axes (Tables 8 and 9); i.e., the region occupied on the first axis by the wet rich woodland, and on the second axis by the Black Spruce forest, as well as part of the wet rich woodland and White Spruce woods (Figure 5). Consistent with this finding was the fact that on the second axis (though not the first), elevation above sea level was negatively correlated (Table 7) at a fair- ly high level (-0.40), and there were similar negative correlations (though smaller) for other elevation-asso- ciated topographic factors (i.e. slope, distance to the coast, and cumulative heat units). The other group of attributes showing a significant relationship with the ordination were “stand origin / management history” and “status in 1935” (both indicative of the past history of a site, particularly human disturbance). These factors showed a high level of association with the second axis (Table 9): stands recorded as “old field’ and as “cleared land in 1935” occurred much more frequently than expected due to chance in the combined third and fourth quar- tiles — the part of the axis where most of the plots of the White Spruce woods lie. By contrast, the first quartile, in which the apparently least human-dis- turbed of all of the ground flora groups occurred (Group 5, of the upland hardwood forest), was posi- tively associated with only “nothing evident” (i.e., no evidence of human disturbance/management) and “burn” — for the latter category, see the footnote of Table 4. The analysis of both the quantitative and qualitative attributes thus indicates that elevation, soil drainage and management history show significant associations with the overall distribution of the plots in the first two dimensions of the ordination space. This suggests that they are likely to play a significant role in deter- mining the ground flora species make-up of the forest- types. The results of the ordination are also consistent with the pattern emerging from the comparison of the properties of the individual TWINSPAN forest-types, as evident in the various comparative tables (Tables | to 6). TABLE 7. Pearson’s correlation coefficients between the quantitatively measured topographic attributes and soil properties (of the humus layer) and the scores of the sam- pling points on the first two axes of the DECORANA ordi- nation. (Degrees of freedom vary between 1114 and 1198.)7 AXIS | AXIS 2 TOPOGRAPHIC ATTRIBUTES Elevation ns -0.40*** Slope -0.07** -0.16*** Distance to coast ns -0.16*** Distance to road 0.068* -0.12*** Corn heat units 0.090** -0).17*** SOIL PROPERTIES pH ns 0.14*** % Carbon -0.09 *** ns Nitrogen ns ns Phosphorus mRIGeet -0.20*** Potassium -().24*** -().21*** Calcium -0.10*** 0.21*** Magnesium -0.08 ** ns +Probabilities: *** - p< 0.001; ** - p< 0.01; * - p< 0.05; ns - not significant. Te eh THE CANADIAN FIELD-NATURALIST Vol. 116 Nn \O ON - It is notable that although there were significant dif- ferences in the soil chemical factors between some of the forest-types (Table 3) (the wet rich woodland especially differed from all of the others), none of the soil factors show much correlation with either of the axes (Table 7). This is perhaps not surprising given the limited variation in soil parent materials on the island. the first 2 value, DECORANA axis, as subdivided into four equal quartiles (see Figure 4). For each attribute those individual classes whose x~ contributed significantly (d.f. = 1) to the total x are listed under the quartiles with which they had a positive association. Quartile 4 Moderately well drained** Imperfectly-drained* Poorly-drained *** Imperfectly-drained** Poorly-drained*** High water table*** Level land*** Swampy*** Discussion For the first time in the botanical history of Prince Edward Island we have a quantitative record of the woodland plant community-types occurring on the island. What is more, this is based on a large objec- tively-selected sample that has been analysed using objective techniques of classification and ordination. We are thus now able to define the forest-types on Prince Edward Island as they exist at the end of the twentieth century: TWINSPAN has led to the recogni- tion of eleven ground flora groups representing five main forest-types: a wet species-rich and base-rich swamp-type woodland; upland hardwood forest; dis- turbed conifer-dominated forest; old field White Spruce woods; and Black Spruce forest. Because of the large amount of data of various kinds collected in this study we are able to describe each of these forest- types in considerable detail, not only in terms of their ground flora, but also their tree components, and the topographic, historical/management and soil factors acting on them. The DECORANA ordination has indicated that in overall terms two environmental factors, soil drainage and forest disturbance, appear to be particularly sig- nificant in determining the forest-types of the island. It is they (and especially drainage indicators) that show a high degree of association with the position of the plots along the first two ordination axes. The importance of these two factors was also borne out in the comparative analysis of the TWINSPAN forest- types: differing soil moisture levels are an important factor distinguishing some of the forest-types (Table 2). At the same time the importance of human-associ- ated forest disturbance and clearance is evident throughout the analysis: all of the TWINSPAN forest- types have a proportion of plots showing evidence of various types of human intervention (e.g. clear-cut- ting, partial cutting, old field status) (Tables 4 and 5), and it is human activity per se which is the most important factor accounting for one of the forest-types (White Spruce woods), and which contributes to a second type, the disturbed conifer-dominated forest. The degree of human-disturbance also appears to dif- | ferentiate the three upland hardwood plot groups. It is not possible to compare these findings in any depth with previous descriptions of the forests of the island, as there are no previous studies of comparable detail. However, it is evident that the general points emerging from this study do not conflict with the brief Quartile 3 Well + moderately drained * > Quartile 2 Well-drained* Quartile | drained*** Cleared *** Rapidly- SS SH Cramer’s V 0.228 0.229 0.326 0.204 0.241 0.141 0.182 x? (d.f.) hice ** (9) ere (9) £19.9%** (3) 46.9*** (3) 197.4*** (12) (18) (3) p<001; *=p-<0.05; AES vas p< 0.001; analysis of the association between selected qualitatively-categorized soil and forest stand properties of the sampling points, and the position of the points along 45 x? (from soil series maps) Soil drainage (from soil pits) Soil drainage Root restriction factors Forest type Stand origin / management history Status in 1935 Slope STAND PROPERTIES: SOIL PROPERTIES: Probabilities: *** TABLE 8. ATTRIBUTE 2002 SOBEY AND GLEN: FORESTS OF PRINCE EDWARD ISLAND 597 TABLE g. x? analysis of the association between selected qualitatively-categorized soil and forest stand properties of the sampling points, and the position of the points along the second DECORANA axis, as subdivided into four equal quartiles (see Figure 4). For each attribute those individual classes whose x? contributed significantly (d.f. = 1) to the total x2 value, are listed under the quartiles with which they had a positive association. ATTRIBUTE x’ (d.f.) Cramer’s V Quartile 1 Quartile 2 Quartiles 3 + 4 SOIL PROPERTIES: Soil drainage 245.3*** 0.269 Well-drained*** Imperfectly-drained*** (from soil pits) (6) Poorly-drained*** Soil drainage 181.8*** 0.284 Well/moderately drained*** Imperfectly-drained** (from soil series maps) (6) Poorly-drained*** Root restriction eee 0.269 High water table*** factors (2) Slope eM Meee 0.287 Sloping ground** Level ground*** (2) STAND PROPERTIES: Forest type SUg te? SO" aaa SH* al (8) HS* Stand origin / 214 Gee ONS * 6 Barn** Partial cut* Old field*** management history (12) Nothing evident*** Swampy*** Boggy*** Status in 1935 ODS? FF 231); Forested *** Cleared *** (2) Probabilities: *** = p < 0.001; ** =p <0.01; *=p< 0.05. and qualitative forest descriptions contained in those regional and national studies that have included the island (i.e., Halliday 1937; Rowe 1959; Loucks 1961). Nor are they in disagreement with the three island- based studies that contain limited qualitative descrip- tions of the internal variation in the island’s forests (Stilgenbauer 1929; Erskine 1960; MacDougall et al. 1988). Generally, all of these previous studies consid- ered soil drainage to be especially important in deter- mining the type of forest occurring in particular areas, even if they only did so by making the general and not wholly accurate comment that the upland areas were occupied by tolerant hardwoods and the lowlands by coniferous forest. However, it was only Erskine (1960) (in the most detailed of these previous descrip- tions) who recognised a wet rich lowland forest corre- sponding to the wet rich woodland of this study. By contrast MacDougall et al. (1988) speculated that the limited differences in the fertility of island soil types appeared to have little effect on tree species composi- tion. All of these previous studies also recognised the very important effects of human disturbance on forest differentiation, though those whose aim was to describe the forest in its hypothetical undisturbed state give it less coverage (Halliday 1937; Rowe 1959; Loucks 1961). This study now provides quantitative data to back up these previous speculations: on an island in which there is a limited range of soil parent materials (most are derived from sandstone and similar rocks (MacDougall et al. 1988)) and of climatic variation, the principal intrinsic factor responsible for the varia- tion in the forest vegetation is variation in soil drainage. Drainage in turn is determined by the inter- action between topographic factors (key factors in this respect are elevation and slope), and the nature, espe- cially the texture, of the soil parent material. In addi- tion, in the areas of poor drainage, there is then an important differentiation (due presumably either to slight differences in the base-richness of the soil par- ent materials or to differences in the chemical richness of lateral drainage water), between the lower pH base- poorer soils with higher C/N ratios supporting Black Spruce forest, and the higher pH more base-rich soils with lower C/N ratios supporting a swamp-type forest in which deciduous trees, especially Red Maple, pre- dominate. Superimposed on any differences caused by drainage are the effects of almost three centuries of human interference, involving the initial forest clear- ance for the creation of farmland (which reached its maximum in the early twentieth century), and the con- tinual exploitation of the remaining forest stands for timber and firewood. In this century especially there has also been the abandonment of much of the poorer farmland (Stilgenbauer 1929; Clark 1959; Glen 1997), resulting in its recolonisation by trees in a process of secondary succession. That the information that has emerged from this study is in agreement with the general observations of the few previous workers who have had an interest in the island’s forests is especially pleasing in view of the small plot size (4 m2) used for the assessment of the ground flora, a size imposed by financial con- straints. (Usually much larger plot sizes [e.g., 200 m2 (Bunce 1982) or 450 m2 (Zelazny et al. 1989)] are used in forest studies.) Even though the 4 m2? plot has been shown to have picked up on average only a third i eek air wn i RT Mia ie te Sept 598 of the ground flora species in the c.1000 m? area around a sampling point (Sobey 1993), this has not prevented statistically-distinctive ground flora groups and forest-types emerging in the multivariate analysis. Furthermore, these forest-types, as well as some of the groups, have proved to be recognisable in the field as distinct vegetation-types. They are also consistent with previous information on the forest vegetation of the island, and at the same time have been amenable to logical explanation in terms of environmental and historical factors. Undoubtedly a larger plot size, if combined (as is usual in such studies) with the omis- sion in the field of those sites that were obviously het- erogeneous, might have led to the production of ground flora groups that were more distinctive and discrete, with for example the plots showing less over- lap in the DECORANA ordination (Figure 4), but this would also have masked an important feature of the island’s forests (or any forest for that matter): forest vegetation is likely to be continuously variable in the composition of both its tree canopy and ground flora. In fact, a larger plot size, without the simultaneous elimination of sample plots containing obviously intermediate vegetation-types, would have led to less distinction between the groups. And since any process of eliminating plots would have inevitably involved questionable subjective decisions, a small plot size has thus had benefits, an additional one being that a much larger number of plots were able to be recorded within the limits imposed by the financial constraints. It must however be stated that it is likely that any forest-type with a limited distribution may not have been picked up at the 1200 sampling points, or at least not in sufficient numbers to constitute a recognisable group in the analysis. With reference to Erskine’s (1960) floristic study of the island, as well as to spe- cial habitat surveys (Taschereau 1974; Anonymous 1982) and to New Brunswick studies (e.g., Zelazny et al. 1989), some forest-types that are likely to consti- tute distinct plant communities on the island (and undoubtedly these would have been more common in the past), but which have not emerged in the analysis are: Cedar woods (only 40 sampling points in all con- tained White Cedar, most of these were classified in the wet rich woodland group); a fern- and herb-rich variant of the upland hardwood forest (containing for example White Ash (Fraxinus americana) or Ironwood (Ostrya virginiana) in the canopy); stream valley woodlands (some with American Elm and Black Ash (Fraxinus nigra)); Pine woods (both White and Red: Pinus strobus, P. resinosa); and Hemlock (Tsuga canadensis) stands (the latter two now very rare). To obtain a sufficiently sized sample of such rare woodland types, we would need to actively search for and then sample such sites in a separate selective inventory. The maps showing the distribution of the sampling sites (Figure 3) give an indication of the spatial distri- THE CANADIAN FIELD-NATURALIST Vol. 116 - bution of the five forest-types on the island. The dis- tinctive spatial patterns evident greatly add to the lim- ited qualitative information available before the 1991 Inventory (e.g., Erskine 1960). However, by making use of another element in the 1990-92 forest inventory (the 1990 aerial photographic survey of the island), the next stage in the study will be to map the total dis- tribution on the island of all of the stands of the five forest-types. Acknowledgments D. Sobey thanks Professor G. McKenna, then Dean of Science at the University of Ulster, for enabling him to take six months leave from University duties in 1992 and 1996 in order to carry out this study; Forestry Canada for covering expenses associated with the 1992 work; and the P.E.I. Department of Agriculture and Forestry for providing financial and other support after 1992. He also thanks H. Baglole of the Institute of Island Studies at the University of P.E.I. for arranging the research associateship which enabled him to use library facilities at the University. We thank T. McCann and J. Boyce of the University of Ulster for computer assistance and B. Rushton of the University of Ulster for reading an early draft of this paper. Literature Cited Anonymous. 1982. Prince Edward Island natural areas sur- vey. Department of Biology, University of Prince Edward Island, Charlottetown, P.E.I. Anonymous. 1992. 1990/1992 Prince Edward Island forest inventory: Summary. Forestry Branch, P.E.I. Department of Energy and Forestry, Charlottetown, P.E.I. Bergeron, J.-F., J.-P. Saucier, A. Robitaille, and D. Robert. 1992. Québec forest ecological program. For- estry Chronicle 68: 53-63. Bowling, C., and V. F. Zelazny. 1992. Forest site classifi- cation in New Brunswick. Forestry Chronicle 68: 3441. Bunce, R. G. H. 1982. A field key for classifying British woodland vegetation. Part 1. Institute of Terrestial Ecology, Cambridge, U.K. Clark, A. H. 1959. Three centuries and the Island. Uni- versity of Toronto Press, Toronto. Erskine, D.S. 1960. The plants of Prince Edward Island. Publication 1088. Canada Department of Agriculture. Gauch, H. G. 1982. Multivariate analysis in community ecology. Cambridge University Press, Cambridge, England. Glen, W. M. 1997. Prince Edward Island 1935/1936 forest cover type mapping. Forestry Division, P.E.I. Department of Agriculture and Forestry, Charlottetown, P.E.I. Haddon, B. D. Editor. 1989. Forest inventory terms in Canada (3rd edition). Forestry Canada, Ottawa. Halliday, W. E. D. 1937. A forest classification for Canada. Canada Department of Mines and Resources, Ottawa. Hill, M. O. 1979a. DECORANA — A FORTRAN program for detrended correspondence analysis and recip- rocal averaging. Ecology and Systematics, Cornell Uni- versity, Ithaca, New York, U.S.A. aii a 2002 Hill, M. O. 1979b. TWINSPAN — A FORTRAN program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes. Ecology and Systematics, Cornell University, Ithaca, New York, U.S.A. Ireland, R. R. 1982. Moss flora of the Maritime Provinces. Publications in Botany number 13, National Museum of Canada, Ottawa. Jones, R. K., G. Pierpont, G. M. Wickware, J. K. Jeglum, R. W. Arnup, and J. M. Bowles. 1983. Field guide to forest ecosystem classification for the Clay Belt, Site Region 3e. Ministry of Natural Resources, Government of Ontario, Canada. Loucks, O. L. 1961. A forest classification for the Mari- time Provinces. Proceedings of the Nova Scotia Institute of Science 25: 85-167. MacDougall, J. L., C. Veer, and F. Wilson. 1988. Soils of Prince Edward Island. Agriculture Canada, Charlotte- town, P.EI. Norusis, M. J. 1990. SPSS Base System User Guide. SPSS Inc., Chicago, U.S.A. Rowe, J. S. 1959. Forest regions of Canada. Forestry Branch, Bulletin 123 [Reprinted 1972 with minor revi- sions as: Publication Number 1300, Canadian Forestry Service, Department of the Environment, Ottawa]. Scott, G. A. J. 1995. Canada’s vegetation — A world per- spective. McGill-Queen’s University Press, Montreal and Kingston. Sims, R. A., W. D. Towill, K. A. Baldwin, and G. M. Wickware. 1989. Field guide to the forest ecosystem SOBEY AND GLEN: FORESTS OF PRINCE EDWARD ISLAND 599 classification for northwestern Ontario. Forestry Canada, Ottawa. Sobey, D. G. 1993. Analysis of the ground flora and other data collected during the 1991 Prince Edward Island for- est inventory. I Floristic analysis. FRDA Report SD-010. Forestry Canada, Charlottetown, P.E.L. Sobey, D. G. 1995. Analysis of the ground flora and other data collected during the 1991 Prince Edward Island Forest inventory. II Plant community analysis. P.E.I. Forestry Division, Department of Agriculture, Fisheries and Forestry, Charlottetown, P.E.I., Canada. Stilgenbauer, F. A. 1929. The geography of Prince Edward Island. Ph.D. dissertation, University of Michigan, Ann Arbor, Michigan. Taschereau, P. M. Editor. 1974. Ecological reserves in the Maritimes. Terminal Report of the Scientific Advisory Panel. C.I. IBP Region 7. Watts, S. B. 1983. Forestry handbook for British Columbia (Fourth edition). University of British Columbia. Zelazny, V. F., T. T. M. Ng, M. G. Hayter, C. L. Bowling, and D. A. Bewick. 1989. Field guide to forest site classi- fication in New Brunswick. Harvey-Harcourt and Fundy Site Regions. Forestry Canada, Fredericton, New Brunswick. Zinck, M. 1998. Roland’s flora of Nova Scotia. Nimbus Publishing and the Nova Scotia Museum, Halifax, Nova Scotia. Received 17 April 2001 Accepted 15 October 2002 600 THE CANADIAN FIELD-NATURALIST Vol. 116 APPENDIX 1. The percentage frequency of the ground flora species in the TWINSPAN ground flora plot groups. (TWINSPAN groups of fewer than 10 species have been omitted.) Only species 2 10% frequency in at least one group are listed. Frequencies printed in bold were significantly greater (x? test, d.f. = 1, p < 0.001) than the frequency for the same species in all 1200 plots — such species are termed “selective” (sensu Bunce 1982) for the plot group.! The species are list- ed in the same order as in the data matrix table generated by the TWINSPAN analysis. SPECIES Alnus incana Aster umbellatus Ilex verticillata Ptilidium pulcherrimum Aster puniceus Dryopteris cristata Viola spp. Epilobium angustifolium Equisetum spp. Equisetum sylvaticum Fragaria virginiana Galium triflorum Ranunculus repens Rhytidiadelphus triquetrus Rubus pubescens Lycopus uniflorus Osmunda cinnamomea Typha latifolia Athyrium filix-femina Cornus sericea Galium palustre Impatiens capensis Mitella nuda Mnium spp. Onoclea sensibilis Ribes lacustre Thalictrum pubescens Gymnocarpium dryopteris Phegopteris connectilis Acer spicatum Taxus canadensis Dryopteris carthusiana Oxalis acetosella Viola cucullata Viola macloskeyi Dennstaedtia punctilobula Lycopodium annotinum Huperzia lucidula Lycopodium obscurum Medeola virginiana Trillium undulatum Viola blanda Aralia nudicaulis Monotropa uniflora Rubus canadensis Aster acuminatus Pyrola elliptica Trientalis borealis Clintonia borealis Coptis trifolia Grass spp. 27 26 Upland Hardwood Forest TWINSPAN GROUND FLORA PLOT GROUPS 8 9 10 Disturbed Conifer- White domindated Spruce Forest Woods 7 #/ 5 5 9 7 6 5 6 ih 5 8 7 i; 19 5) & 12 9 7 12 10 18 10 8 5 5 52 19 ii 8 7 17 8 7 5 60 51 36 47 13 22 14 16 * 19 10 10 33 14 Black All Spruce 1200 Forest plots 10 9 17 (Continued) 2002 SOBEY AND GLEN: FORESTS OF PRINCE EDWARD ISLAND 601 APPENDIX 1. Continued SSS OO EEE TWINSPAN GrRouND FLorRA PLoT Groups 3 5 6 Zz 8 9 SIRS | ta Ma 13. 14 Wet Disturbed Rich Upland Conifer- White Black All Wood- Hardwood domindated Spruce * ‘3 Spruce 1200 SPECIES land Forest Forest Woods Forest plots Meee: 8 4 21 6 @ #4 4% .10 8 07. 6. 498. Moss spp. 5 6 7 6 6 10 48 Maianthemum canadense 32 38 74 a 85 92 56 48 33 10.1 0G 58.8 Sorbus americana 9 15 15 5 12 7 8.3 Hieracium aurantiacum 19 | Polytrichum commune ‘i 20 a3 37 46 20 fa 19.0 Aster spp. 5 14 4.2 Veronica officinalis 13 3.2 Amelanchier spp. 10 15 18 7 17 16 8.3 Cornus canadensis 2h) cas 29 9 61 86 10 83 = 2 ee | i Cypripedium acaule a 2.8 Hylocomium splendens 12 16 7 58 17 6 6.7 Linnaea borealis 11 33 22 8 10.2 Pteridium aquilinum 8 11 19 73 8 49 16.8 Viburnum nudum 20 5 7 62 58 21 67 45 49 26.8 Dicranum polysetum 20 15 19 8 14. 45 10.1 Epigaea repens 11 18 2.2 Melampyrum lineare a 6 8 10 2.5 Carex spp. 44 8 23 15 19 16 23 14 58 48 22.3 Pleurozium schreberi po) 8 13 5 72 69 82 71 75 fz @® 41.8 Ptilium crista-castrensis 7 16 17 18 4.3 Vaccinium angustifolium 15 48 26 25 21 eG 14.8 Vaccinium myrtilloides 12 31 7 17 M4 Si 8.2 Cladina rangiferina 25 1.8 Gaultheria procumbens 6 Y 31 At Gaylussacia baccata 21 810 1.3 Kalmia angustifolia 7 46 58 66 8690 13.8 Ledum groenlandicum 17 ab *37 2.8 Nemopanthus mucronata 21 7 42 Way | 22 8.1 Chamaedaphne calyculata 14 0.6 Rhododendron canadense oO. 3 2.3 Gaultheria hispidula 5 8 41 8 a Smilacina trifolia 58 17 1.7 Sphagnum spp. 29 17 92 Sa 13.0 Cladina stellaris 12 0.7 Number of plots 56." TF 196 155 165 118 135 21 12 2 Mean no. of species/plott 13.7 5.6 7.7 6.9 938° 95 6.5 4.0 13.9 8.9 8.9 8.6 Standard error 0.44 0.34 0.23: 020 2.29 " O27 0.29 0.44 y Ae 0.60 0.38 0.13 Scheffé test? af: abe abe’ abe’) ab? ae* PaRcCe sbc d* af* ae* deg d g* d def - 1. The following additional ground flora species were ‘selective’ for the the various TWINSPAN groups but occurred at less than 10% frequency: Rich Wet Woodland: Ribes hirtellum, Caltha palustris, Eupatorium maculatum, Prunus virgini- ana, Equisetum arvense, Circaea alpina, Ribes glandulosum, Lonicera canadensis, Corylus cornuta, Rosa virginiana. Upland Hardwood Forest: TSP Group 6: Smilacina racemosa, Cornus alternifolia; TSP Group 7: Smilacina racemosa, Monotropa uniflora. Disturbed Conifer-dominated Forest: TSP Group 8: Bazzania trilobata. White Spruce Woods: Myrica pensylvanica. *TWINSPAN Group 11 was a species-poor hardwood-related plot-type and Group 12 was a wet species-rich plot-type transitional between TWINSPAN Groups 3 and 13. +Mean number of species per plot: One-way ANOVA (d.f. = 10): F = 53.0, p < 0.001. Scheffé test: a mean with the same letter is significantly different (p < 0.05) from a mean with the letter followed by an asterisk (e.g. “a” is significantly differ- ent from “a*”’). 602 THE CANADIAN FIELD-NATURALIST Vol. 116 APPENDIX 2. Summary descriptions of the TWINSPAN-derived forest-types (based largely on the data contained in Tables 1 to 6). WET RIcH WOODLAND (TWINSPAN Group 3: 156 plots — 13.8%) A forest-type of frequent occurrence, especially at low elevation sites of varying sizes in the western region, as well as parts of the eastern region of the island (see Figure 1 for region boundaries); it is associated with comparatively less acid but poor- ly-drained soils. The tree canopy contains a mixture of broad-leaves and conifers, with Red Maple (Acer rubrum) predomi- nating, and with low but notable levels of White Cedar (Thuja occidentalis), White Ash (Fraxinus americana), American Elm (Ulmus americana) and Speckled Alder (Alnus incana). It is a species-rich community-type at all levels (in trees, shrubs and ground flora), characterised by 44 selective ground flora species*, an exceptionally large number. Key site factors asso- ciated with the group are low elevation, level land, poor soil drainage, poorly and imperfectly drained soil series, higher pH and base content, lower C/N ratio. Key stand factors: mixed Red Maple-dominated forest, older trees, medium stocking rate. This forest-type appears to be a derivative of the pre-European settlement lowland Red Maple forest (sensu Erskine 1960) — though with considerable effects due to cutting and other disturbances. UPLAND HARDWOOD Forest (TWINSPAN Groups 5 + 6 + 7: 428 plots — 38.0 %) A forest-type of widespead occurrence, well-scattered throughout the central region and in specific areas in the eastern and western regions. The tree canopy is dominated by hardwoods, especially Red Maple (Acer rubrum), with a notable contri- bution by the tolerant hardwood species (Sugar Maple (Acer saccharum), Yellow Birch (Betula alleghaniensis) and American Beech (Fagus grandifolia)). There is also a minority softwood component especially of Balsam Fir (Abies bal- samea), and the presence of Trembling Aspen (Populus tremuloides) and White Birch (Betula papyrifera) is indicative of past and continuing human disturbance. The ground flora is of poor to moderate species-richness, and its three sub-groups are characterised collectively by nineteen different selective ground flora species. Key site factors: moderate to higher ele- vations, sloping ground, well-drained soils, well-drained soil series, lowish to moderate organic matter, varying levels of past forest clearance and cutting, small to moderate woodland size. Key stand factors: hardwood-domination. This forest- type is equivalent to the pre-European settlement tolerant hardwood forest of the uplands (sensu Erskine 1960), though subject to varying levels of past and current human disturbance. DISTURBED CONIFER-DOMINATED FOREST (TWINSPAN Groups 8 + 9: 283 plots — 25.1%) A forest-type of frequent occurrence, especially common in larger woodland areas at moderately low elevations in the east and in the far west. It comprises conifer-dominated forest on soils of varying drainage properties — though not on the wettest. The tree canopy is dominated by softwoods, especially Balsam Fir (Abies balsamea), with White, Black and Red spruce (Picea glauca, P. mariana, P. rubens). The ground flora is of moderate species-richness, with 20 different selective species for the two sub-groups. Key site factors: highish levels of partial-cutting/thinning, highish level of past forest clear- ance, generally good drainage. Key stand factors: softwood-domination, moderate stocking rate. It is difficult to equate this forest-type with any of the climax forest-types of Erskine (1960) (though it may include areas that in the pre-European period were occupied by pine forests). Rather it appears to largely represent a forest-type heavily affected by past and con- tinuing human exploitation. WHITE SPRUCE Woops (TWINSPAN Group 10: 135 plots — 12.0%) A forest-type of common occurrence comprising woodlands of varying sizes on well-drained soils at moderate elevations. It has a distinct distributional bias toward the eastern end of the island. This forest-type has a marked association with land formerly cleared for agriculture. The tree canopy is dominated by a single species, White Spruce (Picea glauca), which is generally youngish, even-aged, and of high density. It is a species-poor community at all levels (i.e. in ground flora, shrubs and trees), characterised by only seven selective ground flora species, most of which are either mosses or weedy aliens. Key site factors: “old field” sites; i.e., formerly cleared land, well and rapidly drained soils, sloping land, lowish organic matter content. Key stand factors: White Spruce-domination, immature trees, low tree age and height, high stocking rate. This forest-type corresponds to the successional “‘old field” White Spruce woods of Erskine (1960). BLACK SPRUCE FoREST (TWINSPAN Groups 13 + 14: 80 plots — 7.1%) A rather uncommon forest-type occurring on areas of varying drainage properties at lower elevations especially in parts of the western and eastern regions. The tree canopy is dominated by conifers, above all Black Spruce (Picea mariana), with some Red Spruce (Picea rubens) and Tamarack (Larix laricina). The ground flora is of moderate species-richness, with 20 different selective species in its two sub-groups. Key site factors: level ground, soils of varying drainage from poor to good, swamp and bog characteristics, low pH, high C/N ratio. Key stand factors: Black Spruce-domination, low stocking rate, low tree height, medium tree age. This forest-type corresponds to the lowland Black Spruce forest of Erskine (1960). The two sub-groups (Groups 13 and 14) appear to represent ecologically-valid variants of this forest-type, the first a wetter forest-type with bog characteristics, the second a dryer forest-type with heath characteristics. *Selective ground flora species (sensu Bunce (1982)) are those whose frequency in any TWINSPAN group was signifi- cantly greater (x? goodness of fit, p < 0.001) than in all of the 1200 inventory plots taken together. Population Trends of Nesting Glaucous-winged Gulls, Larus glaucescens, in the Southern Strait of Georgia, British Columbia TERRY M. SULLIVAN!, STEPHANIE L. HAzitt2*, and Morra J. F. LEMon* '11573 84A Avenue, Delta, British Columbia V4C 285, Canada ? Bird Studies Canada, 5421 Robertson Road, Delta, British Columbia V4K 3N2, Canada Current Address: Department of Zoology and Entomology, University of Queensland, St. Lucia, QLD 4101 Australia. 4Canadian Wildlife Service, 5421 Robertson Road, Delta, British Columbia V4K 3N2 Canada Sullivan, Terry, M., Stephanie L. Hazlitt, and Moira J. F. Lemon. 2002. Population trends of nesting Glaucous-winged Gulls, Larus glaucescens, in the southern Strait-of Georgia, British Columbia. Canadian Field Naturalist 116(4): 603-606. Between 1986 and 1999, the number of nesting Glaucous-winged Gulls (Larus glaucescens) at fourteen breeding colonies declined by 31% (1610 nests) in the southern Strait of Georgia, British Columbia. Declines in the number of nesting pairs at surveyed colonies ranged between 6% and 81% with the smallest change occurring on the largest breeding colony in the study area. We suggest that Glaucous-winged Gull breeding population declines in the southern Strait of Georgia may be due to increased frequency of colony disturbance, in particular increased disturbance by Bald Eagles (Haliaeetus leuco- cephalus). Key Words: Glaucous-winged Gulls, Larus glaucescens, declines, disturbance, British Columbia The breeding population of Glaucous-winged Gulls (Larus glaucescens), like many populations of large gulls, increased in the Strait of Georgia during the 20" century (Drury 1973, 1974; Vermeer and Devito 1989). Glaucous-winged Gulls in this region increased from 6150 nesting pairs in 1960 to 13002 pairs in 1986 (Verbeek 1986; Vermeer and Devito 1989). This increase was thought to be the result of increased food abundance as this period correspond- ed with the increased utilization of landfill sites (Drent and Guiguet 1961; Vermeer 1963; Vermeer 1982; Butler et al. 1980). The breeding population of Bald Eagles (Haliae- etus leucocephalus) also increased in the Strait of Georgia from 45 pairs in the 1970s to 97 pairs in 1987 (Trenholme and Campbell 1975; Vermeer et al. 1989). Bald Eagles prey on nesting Glaucous- winged Gulls (Butler et al. 1980; Vermeer et al. 1989: Vermeer and Devito 1989) and flush them from their nests allowing gulls and crows to depre- date eggs and young (Drent et al. 1964; Verbeek 1982). In this paper we document recent changes in the number of nesting pairs of Glaucous-winged Gulls at fourteen breeding colonies. We suggest that the recent decline in numbers of breeding Glaucous- winged Gulls in the southern Strait of Georgia is related to an increased frequency of disturbance at breeding colonies, and in particular disturbance by Bald Eagles. Methods Fourteen Glaucous-winged Gull colonies were censused in 1997 and 1999 (four colonies only in 1997; seven colonies only in 1999; and three small colonies in both years) in the southern Strait of Georgia, approximately 65 km south of Vancouver, British Columbia (49° 15’ N, 123° 07’ W) (Figure 1). The 14 colonies surveyed were selected because the sites coincided with other research initiatives; however, they are only a subset of the 76 islands and small islets used as nesting sites by Glaucous- winged Gulls in the Strait of Georgia (Vermeer and Devito 1989). Between | and 9 July 1997 and 1 and 25 June 1999 each colony was surveyed by one to four observers, depending on size of the colony, fol- lowing survey methods outlined in Vermeer and Devito (1989). Briefly, colonies were censused by walking through the colony while counting nests. In large colonies we placed a small colored jellybean in each nest to ensure we did not re-count nests. The daily presence or absence of Bald Eagles was recorded on Mandarte Island between | May and 31 July 1997. The presence of Bald Eagles was only recorded if we observed a disturbance of nesting gulls on the island. This differs from other studies, which also included Bald Eagles flying near Mandarte Island (Verbeek 1982). Results and Discussion A total of 3506 Glaucous-winged Gull nests were found at 14 colonies, an overall reduction of 1610 nests (31%) from 1986 (Table 1). The number of nesting pairs increased at a single colony, Grebe Islet, while nesting pair numbers dramatically decreased at the other 13 colonies. Reduction in nesting pairs ranged from 6% on Mandarte Island to 81% on Imrie Island. As a result, the distribution of 603 604 87 Snake Island .* Five Finger lsand Hudson Rocks “K Pam Rocks Christie Islet Arbutus Isand Imrie Island Greig Islets Little Group Islands Mandarte Island Mandarte South Islet Sallas Rocks kilometres THE CANADIAN FIELD-NATURALIST Vol. 116 FiGurRE 1. Location of the 14 Glaucous-winged Gull breeding colonies surveyed in the south- ern Strait of Georgia, British Columbia. nesting Glaucous-winged Gulls shifted such that 61% of the nesting gulls were found on the largest breeding colony, Mandarte Island, in 1997 compared to 45% in 1986 (Table 1). This is the first reported decline in the number of nesting Glaucous-winged Gulls within the Strait of Georgia in almost four decades. The decline in the number of nesting pairs of Glaucous-winged Gulls could be related to an increase in colony disturbances, and in particular increased disturbance by Bald Eagles. On Mandarte Island, Bald Eagles were observed flushing gulls on all days researchers were present on the island in 1997 (Table 2). In addition, the number of eagle dis- turbances of nesting gulls within each day, in 1997, ranged from | to 23 times, with up to six eagles on Mandarte Island at any one time. Colony distur- bances during the nesting period result in egg and chick predation by Northwestern Crows and other Glaucous-winged Gulls (Verbeek 1982). On Mandarte Island, over 50 depredated gull eggs were found in a single crow cache in both 1996 and 1997. In addition to the increased number of breeding Bald Eagles in the Strait of Georgia, more than twice the numbers of eagles were observed in 1987 when compared to 1974 (Vermeer et al. 1989). An increase in the observed number of eagles on or near Mandarte Island was not detected until the late 1970s and early 80s. Since that time, eagle observa- tions occurred almost every day during the gull breeding season. Bald Eagles have negatively impacted other colo- nial nesting species in the Strait of Georgia. Increased Bald Eagle predation resulted in decreased breeding productivity of Great Blue Herons (Ardea herodias fannini) (Butler 1997; Vennesland 2000) and disturbances by Bald Eagles at Double-crested Cormorant colonies have caused delays in the onset of incubation by up to a month and a half (Sullivan 1998). However, the number of breeding Glaucous- winged Gulls only declined by 6% on Mandarte Island, a site with a high degree of Bald Eagle activity. There are other possible sources of disturbance at gull colonies, including the presence of River Otters (Lutra canadensis) and humans. Otter trails and scat were found at all colonies during our surveys. River Otters prey upon Glaucous-winged Gulls (Foottit and Butler 1977; Verbeek and Morgan 1978) and were responsible for reproductive failure on Imrie Island in 1985 (Vermeer and Devito 1989). However, there is no current information on the 2002 SULLIVAN, HAZLITT, AND LEMON: NESTING GLAUCOUS-WINGED GULLS 605 Table 1. Comparisons of the number of Glaucous-winged Gull nests from 1986 to 1997 or 1999 in the southern Strait of Georgia, British Columbia. —.)) SS hI TT % Change from 1986 to GWGU Colony 1986! 1997 1999 1999 (or 1997) Arbutus Island 150 97 55 —63 Imrie Island 216 33 41 -81 Greig Islets 52 5 14 —73 Little Group Islands 6 3 ns —50 Mandarte Island 2259 2124 ns —6 Mandarte South Islet 104 97 ns —7 Sallas Rocks 29 13 ns —55 Snake Island 673 ns 204 —70 Five Finger Island 671 ns 288* —57 Hudson Rocks 247 ns 122 —51 Pam Rocks 109 ns 25 —77 Christie Islet 454 ns 232* —49 Bird Islet 38 ns 31 -18 Grebe Islets 108 ns af +138 Total 5116 3506** -31 '1986 data from Vermeer et al. (1989); ns = not surveyed. *Only 80% of the gull colony counted to avoid disturbance of nesting cormorants. Number estimated based on coverage of the colony. **Sum of 1999 counts at 10 colonies and 1997 counts for 4 colonies with 1997 data only. status of River Otter populations in the Strait of Georgia. tat changes rather than to increased raptor popula- tions. Human activities can also affect the number of disturbances at nesting colonies. Campfire pits and tent sites were found on many of the islands with nesting gulls. In addition, Verbeek (1982) detected an increase in the number of Bald Eagles flying over Mandarte Island on weekends and suggested that this was due to roosting eagles being disturbed by boaters at nearby islands. Recent decreases in the breeding abundance of other large gulls in the northern hemisphere have been documented, including declines in breeding Herring Gulls (Larus argentatus) and Glaucous Gulls (Larus hyperboreus) (Gilchrist and Robertson 1999: Robertson et al. 2001). These declines have been attributed to decreased food supplies and habi- We conclude that there has been an overall decline in the number of nesting Glaucous-winged Gulls at 14 colonies in the southern Strait of Georgia. We suggest that the observed decline in the numbers of breeding Glaucous-winged Gull pairs may be due to an increased frequency of disturbance by Bald Eagles, although there are many possible alternative explanations. We strongly recommend continued monitoring of the breeding Glaucous-winged Gull population in the Strait of Georgia. Acknowledgments We thank J. Cotter, L. Bosschieter, J. G. Burns, N. Wolfe and S. Nebel for their help with the fieldwork. We thank R. Butler for his comments on previous Table 2. The number of days observers were present on Mandarte Island (# Days) and the number of days Bald Eagles were observed on or flying by (Eagle Days) Mandarte Island, British Columbia. May June July Total Year #Days Eagle Days (%) # Days Eagle Days(%) #Days Eagle Days (%) # Days Eagle Days (%) 1959! 31 13 (41) 30 6 (20) 31 2 (6) 9] 21 (23) 1960! 31 12 (38) 30 7 (23) 31 4 (13) 91 23 (25) 1976! 31 14 (45) 30 6 (20) 29 2 (7) 90 22 (24) 1977! 31 17 (55) 30 14 (47) 29 12 (41) 90 43 (48) 1978! 31 14 (45) 30 9 (30) 31 7 (23) 9] 30 (33) 1979! 31 20 (65) 30 21 (70) 22 5 (23) 83 46 (55) 1980! 31 21 (68) 30 24 (80) 21 15 (71) 82 60 (73) 1989 15 10 (66) 20 20 (100) 16 12 (75) 51 42 (82) 1996 30 30 (100) 30 30 (100) 22 17 (77) 82 77 (94) 1997 16 16 (100) 21 21 (100) 22 22 (100) 59 59 (100) 'Data from Verbeek (1982). 606 drafts of this paper. Financial support for M. Lemon was provided by the Canadian Wildlife Service, Pacific and Yukon Region. Financial support for S. L. Hazlitt was provided by the Canadian Wildlife Service, Pacific and Yukon Region, the Center for Wildlife Ecology Research at Simon Fraser Univer- sity, and the John K. Cooper Foundation. 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Simon Fraser Univer- sity, Burnaby, British Columbia. Verbeek, N. A. M. 1982. Egg predation by Northwestern Crows: its association with human and Bald Eagle activity. Auk 99: 347-352. Verbeek, N. A. M. 1986. Aspects of the breeding biology of an expanded population of Glaucous-winged Gulls in British Columbia. Journal of Field Ornithology 57: 22-33. Verbeek, N. A. M., and J. L. Morgan. 1978. River Otter predation on Glaucous-winged Gulls on Mandarte Island, British Columbia. Murrelet 59: 92-95. Vermeer, K. 1963. The breeding ecology of the Glaucous- winged Gull (Larus glaucescens) on Mandarte Island, B.C. Occasional Paper 13, British Columbia Provincial Museum, Victoria, B.C. 104 pages. Vermeer, K., and K. Devito. 1989. Population trend of nesting Glaucous-winged Gulls in the Strait of Georgia. In The ecology and status of marine and shoreline birds in the Strait of Georgia, British Columbia. Edited by K. Vermeer and R. W. Butler. Special Publication, Can- adian Wildlife Service, Ottawa, Ontario. Vermeer, K., K. H. Morgan, R. W. Butler, and G. E. J. Smith. 1989. Population, nesting habitat and food of Bald Eagles in the Gulf Islands. Jn The ecology and sta- tus of marine and shoreline birds in the Strait of Georgia, British Columbia. Edited by K. Vermeer and R. W. Butler. Special Publication, Canadian Wildlife Service, Ottawa. Received 3 May 2001 Accepted 18 September 2002 Extralimital Occurrences of Ringed Seals, Phoca hispida, on Sable Island, Nova Scotia ZOE N. Lucas! and DONALD F. MCALPINE? 'P.O Box 64, Halifax CRO, Halifax, Nova Scotia B3J 2L4 Canada *New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada; dmcalpin@nb.aibn.com Lucas, Zoe N., and Donald F. McAlpine. 2002. Extralimital occurrences of Ringed Seals, Phoca hispida, on Sable Island, Nova Scotia. Canadian Field-Naturalist 116(4): 607-610. First records for the Ringed Seal, Phoca hispida, in Nova Scotia are documented. Most of these records are from Sable Island. These observations coincide with documented increases in extralimital records of other arctic seal species in the region. However, occurrences of vagrant Ringed Seals may not be due to the same factors responsible for recent increases in extralimital observations of Harp (Pagophilus groenlandica) and Hooded (Cystophora cristata) Seals in the Northwest Atlantic. Key words: Distribution, Ringed Seal, Phoca hispida, Sable Island, Nova Scotia. In the Northwest Atlantic during the 1990s there was a dramatic increase in the numbers of Harp (Pagophilus groenlandica) and Hooded (Cystophora cristata) seals observed outside their historic north- ern range (McAlpine and Walker 1990, 1999; Stevick and Fernald 1998; McAlpine et al. 1999a,b). This increase was observed on Sable Island, a 40 km long sand bar located about 250 km south-east of Halifax, Nova Scotia (Lucas and Daoust 2002). During this same period Ringed Seals, Phoca hispi- da, were recorded on the island for the first time. These observations provide the first confirmed records of the Ringed Seal for Nova Scotia. Here we document and comment on these observations. The Ringed Seal is the most abundant seal in northern waters and is circumpolar in distribution. The species is widely distributed throughout the Canadian Arctic, in eastern Canadian marine waters occurring from Newfoundland and Labrador north- ward to the North Pole (Kingsley 1990; Reeves 1998). Peterson (1966) notes that this seal will some- times venture south of the Strait of Belle Isle as far as La Tabitiére, Quebec and South Brooks, Newfoundland, and Hannah (2000) reports that the Ringed Seal will move southward with the formation of ice down the north Shore of the Gulf of St. Lawrence as far as Harrington Harbour, Quebec. Both Peterson (1966) and Banfield (1974) plot an 1888 record from Manicaugau, well up the mouth of the St. Lawrence, as extralimital. Ringed Seals occupy areas of stable, dense, sea-ice for most of the year. Although preferred breeding habitat appears to be land-fast ice in fiords and bays, this species is also widely distributed in drifting, off- shore, pack ice, where some breeding has been observed (Reeves 1998; Wiig et al. 1999). Ringed Seals can remain in the Arctic year-round because of their ability to maintain breathing holes in very thick ice (Kingsley 1990). Although it has been assumed that adult Ringed Seals are largely sedentary (McLaren 1958), with young-of-the-year dispersing over distances up to several hundred kilometers, tag- ging studies in Greenland (Kapel et al. 1998) and in Arctic Canada (Smith 1987) have demonstrated dis- persal distances for sub-adult Ringed Seals of more than 1000 km from site of release. Ridoux et al. (1998) note that extreme extralimital occurrences of arctic seals are often considered to be those of young and inexperienced animals. However, their observa- tions suggest that juveniles have sufficient navigation- al skills to return to their usual habitat after traveling long distances. A juvenile Ringed Seal tagged and released at the western end of the English Channel in March 1995 was found among the stomach contents of a Greenland Shark (Somniosus microcephalus) in May 1996 north of Iceland. Assumming the seal was ingested by the shark in waters near Iceland, the Ringed Seal had traveled a minimum straight-line dis- tance of 2600 km in five months (Ridoux et al. 1998) There are no previously published reports of the Ringed Seal from Nova Scotia (Scott 1968) and cur- rently no records of the Ringed Seal from New Brunswick. However, three Ringed Seals were reported from mainland Nova Scotia between 1991-1994 (T.Wimmer, personal communication to ZNL). Unfortunately, only minimal details for these mainland Nova Scotia records are available. One Ringed Seal was found at the mouth of the Sackville River, Bedford, in 1993. A second Ringed Seal, fat and healthy, but with several cuts on the body, was found at the same location on 2 March 1994. We have been unable to locate details on the third mainland Nova Scotia Ringed Seal. There are also recent observations from the eastern seaboard, exclusive of Nova Scotia, of Ringed Seals well south of their normal range. From 1989-2002 approxi- 607 608 mately 15 Ringed Seals have been recorded from Maine to New Jersey (B. L. Rubinstein, personal communication to ZNL), including two immature females found in New Jersey in late January 1990: one dead and one live, the latter approximately 85 cm in length (B. Schoelkopf, personal communi- cation to ZNL). Fourteen Ringed Seals have been reported from Sable Island since 1990 (Table 1). Identification of Ringed Seals was based on a pelage pattern showing distinct light-coloured rings on the dorsal surface of an otherwise dark-grey pelt, on relatively small body size and delicate head shape, and on comparison of skeletal features from seven of the animals (NBM 5851,5921,5932,5947,5948,5949,5950) with refer- ence material in the New Brunswick Museum collec- tion. In one specimen (NBM 5921) the characteristic pelage pattern was largely absent. Some light- coloured ring markings were present on the flanks, but otherwise the dorsal surface was a uniform grey. Measured length is available for only six of the specimens found on Sable Island during 1990-2002; all were under 100 cm (Table 1). Ringed Seal pups in the eastern Canadian Arctic are born during early to mid-April at about 65 cm in length (Smith and Stirling 1975; McLaren 1993), and are suckled for about two months. Based on growth rates given in McLaren (1993), age of the six known-length speci- mens was probably 10 months for two, and likely 10 to 22 months for the other four. The carcass of a Table 1. Ringed Seals observed on Sable Island 1990-2002. THE CANADIAN FIELD-NATURALIST nnn Cn Ln NC NN Me i Vol. 116 - shark-killed seal (Figure 1), though very fresh, was extensively damaged and it was only possible to estimate length at roughly 100 cm or less. The remaining seven seals were less than 100 cm in length. Thus, all 14 Ringed Seals found from 1990-2002 were young animals. Gut contents were examined in five specimens. The stomach of the female found 24 February 1998 contained a small amount of sand. The stomach of the male found on March 11 1998 contained a small amount of sand, one squid beak and two fragments of plastic, with no parasites and the colon was packed with a mix of dry solid feces and sand. The stomachs of the 14 December 1998, 20 January 2002, and 1 July 2002 animals were empty. King (1983) sug- gested that Ringed Seals may use suction in feeding, and thus may ingest sand when feeding near the bot- tom. The stomach of the January 1998 female killed at sea by a shark was empty, but the colon contained feces including many small unidentified otoliths. Of the eight Ringed Seals for which blubber measurements are available, seven intact beached carcasses had mean dorsal and sternal blubber thick- nesses of 9.1 mm (SD = 3.1 mm dorsal, SD = 3.9 mm sternal), whereas the one shark-killed Ringed Seal had dorsal and sternal blubber thickness of 45 mm and 30 mm, respectively. The sternal blubber thickness of the shark-killed Ringed Seal was within the range for the one- to four-year-old seals collected by McLaren (1958) in southwest Baffin Island and Date Sex vo iy G DB SB Observer Comments January 1990 m — — — — Skip J. F. Young live, healthy, January 1992 — — — — — G. Forbes, B. Beck 2 live, active; 1 on north, 1 on south side of island. February 1995 — os — — — Z. Lucas live, active. 14 March 1996 m — —- —- — Z. Lucas live, small cuts on hind flippers. 14 January 1997 fi -— — 15 10 Z. Lucas found dead (NBM 5947). 25 January 1997 m 79 35 6 8 Z. Lucas found dead (NBM 5948). 24 February 1997 f 67 47 5 6 Z. Lucas live, died 4 days later (NBM 5949) 11 April 1997 m 72 49 5 6 Z. Lucas live, died 1 day later (NBM 5851) 5 January 1998 f — 80 45 30 Z. Lucas shark kill, Figure 1. 12 March 1998 m a — — — Z. Lucas live, very thin 14 December 1998 m 95 60 13 ES Z. Lucas dead (NBM 5950) 20 January 2002 f 69 52 12 1] Z. Lucas fresh dead (NBM 5921) 1 July 2002 m ~80 62 8 10 Z. Lucas found dead, scavenged (NBM 5932) 'SL= standard length, G= girth, DB= dorsal blubber thickness, SB= sternal blubber thickness. SL and G recorded in cm; DB and SB recorded in mm. 2002 1998 (Z. Lucas photo). northwest Foxe Basin during March to September. The shark-killed Ringed Seal was in better condition than the seals that died on the beach. Lucas and Daoust (2002) found that the dorsal and sternal regions of the blubber in intact carcasses of Harp and Hooded seals found on the beach were significantly thinner than in shark-killed animals. This indicates that while Harp and Hooded seals which come ashore are likely to be those animals in poor condi- tion (McAlpine and Walker 1990), those that are killed by sharks may represent seals which are healthier and feeding at sea, and less likely to haul out. Thus, the greater blubber thickness and the pres- ence of fish otoliths in the gut of the shark-killed Ringed Seal suggests that healthy individuals of this species may be found in waters of the Scotian Shelf. While the dramatic increase in numbers of Harp and Hooded seals on Sable Island is well-document- ed (Lucas and Daoust 2002), the apparent increase in sightings of Ringed Seals may, in part, be the result of beach surveys for Harp and Hooded Seals carried out from 1995 to 1998. Eight of 14 Ringed Seals were found during these surveys, which were discon- tinued after spring 1998. Prior to these systematic surveys for Harp and Hooded seals, Ringed Seals may have been overlooked among the large numbers of juvenile Grey (Halichoerus grypus) and Harbour (Phoca vitulina) seals hauled out on Sable Island (Boulva and McLaren 1979; Mansfield and Beck 1977; Stobo and Zwanenburg 1990). Although Fisheries and Oceans Canada have car- ried out research on seals on Sable Island during January-February and May—June of most years since 1965, the focus of these studies has been Grey and LUCAS AND MCALPINE: RINGED SEALS ON SABLE ISLAND ti Si» fer Oe Jes agg <¥, is co a a ws « cae 7 = — FiGuRE 1. Juvenile female Ringed Seal, shark-killed, Sable Island, Nova Scotia, 5 January 609 - C3 ed: 3 o”™.. Harbour seals. If Ringed Seals were observed, these sightings were not recorded. Additionally, Ringed Seals resemble juvenile Harbour Seals in locomotion on land and superficially in appearance. Van Bree (1997) notes that even trained zoologists may have difficulty correctly identifying Ringed Seals. He suggests that increases in reports of arctic seals, including the Ringed Seal, on the western European continental coast should be attributed to previous misidentification, an increase in the number of natu- ralists, better identification manuals, and the creation of seal rehabilitation centers. While the extralimital occurrences of Ringed Seals reported here coincide with a dramatic increase in numbers of Harp and Hooded seals on the Island, extralimital occurrences of Ringed Seals may not be due to the same factors. Reasons for the increase in numbers of extralimital occurrences of the more migratory Harp and Hooded seals remain speculative (McAlpine et al. 1999a; Lucas and Daoust 2002), but increases in Harp Seals have been linked to changes in environmental conditions (Lacoste and Stenson 2000). Acknowledgments We are grateful to G. Forbes and staff of the Sable Island Station for logistical advice and assistance, and to T. Wimmer, Nova Scotia Stranding Network, Halifax, for providing details on mainland Nova Scotia records of Ringed Seals from the files of the late J. Parsons. B. L. Rubinstein, New England Aquarium, B. Schoelkopf and staff of the Marine Mammal Stranding Center, Brigantine, New Jersey, and S.J. F. Young, Vancouver Aquarium Marine 610 Science Centre, all generously shared information with us. The following institutions also provided assistance: the Nova Scotia Petroleum Directorate, EnCana Resources, and ExxonMobil Canada. Literature Cited Banfield, A. W. F. 1974. The mammals of Canada. University of Toronto Press, Toronto. Boulva, J., and I. A. McLaren. 1979. Biology of the har- bour seal, Phoca vitulina, in eastern Canada. Bulletin of the Fisheries Research Board of Canada 200: 1-24. Kapel, F. O., J. Christiansen, M. P. Heide-Jgrgensen, T. Harkonen, E. W. Born, L. @. Knutsen, F. Riget, and J. Teilmann. 1998. Netting and conventional tagging used to study movements of ringed seals (Phoca hispida) in Greenland. Pages 211-228 in Ringed Seals in the North Atlantic. Edited by Mads Peter Heide- Jgrgensen and Christian Lydersen. NAMMCO Scientific Publications, Volume 1, The North Atlantic Marine Mammal Commission, Troms¢, Norway. Hannah, J. 2000. Seals of Atlantic Canada and the North- eastern United States. International Marine Mammal Association Inc., Guelph, Ontario. King, J. E. 1983. Seals of the World. Second edition. British Museum of Natural History and Oxford Uni- versity Press, London. Kingsley, M. C. S. 1990. Status of the Ringed Seal, Phoca hispida, in Canada. Canadian Field-Naturalist 104: 138-145. Lacoste, K. N., and G. B. Stenson. 2000. Winter distribu- tion of harp seals (Phoca groenlandica) off eastern Newfoundland and southern Labrador. Polar Biology 23: 805-811. Lucas, Z. N., and P.-Y. Daoust. 2002. Large increase of Harp Seals (Phoca groenlandica) and Hooded Seals (Cystophora cristata) on Sable Island, Nova Scotia, since 1995. Polar Biology 25: 562-568. Mansfield, A. W., and B. Beck. 1977. The Grey Seal in eastern Canada. Fisheries and Marine Service Technical Report 704: 1-81. McAlpine, D. F., P. T. Stevick, and L. D. Murison. 1999a. Increase in extralimital occurrences of ice-breed- ing seals in the Northern Gulf of Maine Region: More seals or fewer fish? Marine Mammal Science 15: 906-911. McAlpine, D. F., P. T. Stevick, L. D. Murison, and S. D. Turnbull. 1999b. Extralimital records of Hooded Seals (Cystophora cristata) from the Bay of Fundy and north- ern Gulf of Maine. Northeastern Naturalist 6: 225-230. McAlpine, D. F., and R. J. Walker. 1990. Extralimital records of the Harp Seal, Phoca groenlandica, from the THE CANADIAN FIELD-NATURALIST Vol. 116 - western North Atlantic: a review. Marine Mammal Science 6: 248-252. McAlpine, D. F., and R. J. Walker. 1999. Additional records of the Harp Seal, Phoca groenlandica, from the Bay of Fundy, New Brunswick. Canadian Field- Naturalist 113: 290-292. McLaren, I. A. 1958. The biology of the Ringed Seal (Phoca hispida Schreber) in the eastern Canadian Arctic. Bulletin of the Fisheries Research Board of Canada 118: 97 pages. McLaren, I. A. 1993. Growth in pinmipeds. Biological Reviews 68: 1-79. Peterson, R. L. 1966. The mammals of eastern Canada. Oxford University Press, Toronto. Reeves, R. 1998. Distribution, abundance and biology of Ringed Seals (Phoca hispida): an overview. Pages 9-45 in Ringed Seals in the North Atlantic. Edited by Mads Peter Heide-Jgrgensen and Christian Lydersen. NAMM- CO Scientific Publications, Volume 1, The North Atlantic Marine Mammal Commission, Tromsg, Norway. Ridoux, V., A. J. Hall, G. Steingrimsson, and G. Olafsson. 1998. An inadvertent homing experiment with a young Ringed Seal, Phoca hispida. Marine Mammal Science 14: 883-888. Scott, F. C. 1968. The seals of Nova Scotian waters. Nova Scotia Museum, Halifax, Nova Scotia. 10 pages. Smith, T. G., and I. Stirling. 1975. The breeding habitat of the Ringed Seal (Phoca hispida): The birth lair and associated structures. Canadian Journal of Zoology 52: 1297-1305. Smith, T. G. 1987. The Ringed Seal, Phoca hispida, of the Canadian western Arctic. Canadian Bulletin of Fisheries and Aquatic Sciences 216: 1-81. Stevick, P. T., and T. W. Fernald. 1998. Increase in extralimital records of Harp Seals in Maine. Northeastern Naturalist 5: 75-82. Stobo, W. T., and K. C. T. Zwanenburg. 1990. Grey seal (Halichoerus grypus) pup production on Sable Island and estimates of recent production in the north- west Atlantic. Canadian Bulletin of Fisheries and Aquatic Sciences 222: 171-184. Van Bree, P. J. H. 1997. On extralimital records of arctic seals (Mammalia, Pinnipedia) on the western European continental coast in the past and present — a summary. Beaufortia 47: 153-156. Wiig, @., A. E. Derocher, and S. E. Belikov. 1999. Ringed Seal (Phoca hispida) breeding in the drifting pack-ice of the Barents Sea. Marine Mammal Science 15: 595-598. Received 7 May 2001 Accepted 24 December 2002 Chromosome Numbers Determined from Canadian and Alaskan Material of Native and Naturalized Mustards, Brassicaceae (Cruciferae) GERALD A. MULLIGAN Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada Mulligan, Gerald A. 2002. Chromosome numbers determined from Canadian and Alaskan material of native and natural- ized mustards, Brassicaceae (Cruciferae). Canadian Field-Naturalist 116(4): 611-622. All herbarium specimens documenting chromosome numbers of 154 species and varieties of Canadian and Alaskan Brassicaceae (Cruciferae) were examined and their identity and nomenclature standardized. Included for each are the chro- mosome numbers, areas where the counted material was collected, publication references, voucher collectors and collec- tion numbers, and the names of the herbarium repositories. All specimen-vouchered published records and a small number of previously unpublished chromosome number determinations are included. Key Words: Mustards, Brassicaceae, Cruciferae, chromosome numbers, Canada, Alaska Many chromosome numbers have been deter- mined from Canadian and Alaskan material of native or naturalized mustards, Brassicaceae (Cruciferae), and the results published. These numbers were obtained either from material collected in nature or from plants cultivated from seed or roots taken from natural habitats. Voucher specimens were deposited in herbaria by most authors. Since different taxo- nomic treatments were used to identify the plants studied or, in a few instances, misidentifications occurred, I am standardizing the names of the taxa according to the taxonomy used in my recent Key to the Brassicaceae (Cruciferae) of Canada and Alaska (Mulligan 2002)*. I have been successful in obtain- ing for study virtually all of the chromosome vouch- ers. The most important exceptions are the herbari- um vouchers for the many chromosome numbers published for Manitoba Brassicaeae by Askell and Doris Live (Love and Love 1975, 1982; Léve and Ritchie 1966). I have not seen annotated voucher specimens for any of their published chromosome numbers of Manitoba Brassicaceae. The authors stat- ed that they had deposited vouchers, in either WIN or COLO. There are only a few unannotated, possi- ble vouchers, in WIN and no Loéve specimens in COLO. There are also no Léve Brassicaceae chro- mosome vouchers in CAN, DAO, MTJB, or MT, other possible repositories. Since the identifications cannot be checked and standardized without vouch- ers, I have, reluctantly, excluded most of their pub- lished chromosome numbers of Manitoba Brassicaceae from the following listing. For unifor- mity and to conserve space, all haploid numbers (n) are listed as somatic numbers (2n). Literature references for any additional chromo- *see Documents Cited 611 some numbers, determined on Canadian or Alaskan material of native or naturalized Brassicaceae, would be appreciated. These can be sent to me at the above address or e-mailed to mulligan4520@rogers.com. Geographical Abbreviations AK: Alaska YT: Yukon NT-M: Northwest Territories, Mackenzie District N-F: Nunavut, Keewatin District NF: Newfoundland and Labrador PE: Prince Edward Island NS: Nova Scotia NB: New Brunswick PQ: Québec ON: Ontario MB: Manitoba SK: Saskatchewan AB: Alberta BC: British Columbia Herbaria Cited Abbreviations ALA: University of Alaska, Fairbanks, AK ALTA: Botany Department, University of Alberta, AB CAN: Botany Division, National Museum of Natural Sciences, Ottawa, ON [= Canadian Museum of Nature]. COLO: University of Colorado Museum, Boulder, Colorado DAO: Vascular Plant Herbarium, Agriculture and Agri- Food Canada, Ottawa, ON DS: Stanford University, San Francisco, California GH: Botany Department, Harvard University, Cambridge, Massachusetts HAM: Royal Botanical Gardens, Hamilton, ON MT: Institut botanique, Université de Montréal, PQ MTJB: Jardin botanique de Montréal, Montréal, PQ QFA: Faculté des Sciences de l’agriculture et de l’alimenta- tion, Université Laval, Sainte-Foy, PQ 612 THE CANADIAN FIELD-NATURALIST Vol. 116 QUE: Herbier du Québec, Complex scientifique, Sainte- | UPS: Botanical Museum, Uppsala University, Uppsala, Foy, PQ. Sweden S: Botany Department, Swedish Museum of Natural WIN: Botany Department, University of Manitoba, History, Stockholm Winnipeg, MB UBC: Botany Department, University of British Columbia, Vancouver, BC Systematic List Alliaria petiolata (M. Bieb.) Cavara & Grande 2n = 42, ON, Grainger & MacLeod 4155 (DAO), Mulligan (1984) as Alliaria officinalis. Alyssum alyssoides (L.) L. 2n = 32, BC, Mulligan & Mosquin 2684 (DAO), Mulligan (1964). Alyssum desertorum Stapf 2n = 32, AB, Mulligan & Mosquin 2684 (DAO), Mulligan (1964). Alyssum obovatum (C. A. Mey.) Turcz. (= Alyssum americanum Greene) 2n = 30, AK, Batten & Dawe 78-392 (ALA), Dawe & Murray (1981a) as Alyssum americanum. Arabidopsis salsuginea (Pallas) O. E. Schulz (= Thellungiella salsuginea (Pallas) N. Busch) 2n = 14, YT, Calder & Gillett 25786 & 26489 (DAO) and Calder 28344 (DAO), all in Mulligan (1964) as Thellungiella salsuginea. Arabidopsis thaliana (L.) Heynh. 2n = 10, ON, Grainger 4005 (DAO), Mulligan (1984). Arabis alpina L. 2n = 16, PQ, Knowlton (GH), Rollins (1941); Cing-Mars et al., 6 juil., 1960 (DAO), Mulligan (1964). Arabis arenicola (Richardson ex Hook.) Gelert 2n = 16, PQ, Hedberg 3408b (UPS), Hedberg (1967); Lepage 39394 (DAO), Mosquin in Mulligan (1995). Arabis caucasica Willd. (= Arabis alpina subsp. caucasica (Willd.) Briq.) 2n = 16, ON, Mulligan 2829 (DAO), Mulligan (1964) as Arabis albida. Arabis columbiana Macoun 2n = 14, BC, Mosquin & Mulligan 4930 & 4937 (DAO), Mosquin in Mulligan (1995). 2n = 20+2B, BC, Calder & Spicer 33480 (DAO), Mulligan (1964) as Arabis divaricarpa. Arabis divaricarpa var. dacotica (Greene) B. Boivin 2n = 14, MB, Love & Love 5373 (DAO, WIN), Love & Love (1975) as Boechera divaricarpa. 2n = 21, YT, Calder & Kukkonen 28067 (DAO), Mulligan (1964). BC, Calder & Gillett 26501 & 26648 (DAO), Mulligan (1964). 2n = 28, NT-M, Cody & Spicer 11451 (DAO), Mulligan (1964). MB, Mosquin & Mosquin 4904 & 4910 (DAO), Mosquin in Mulligan (1995). BC, Calder & Kukkonen 26749 (DAO), Mulligan (1964). Arabis divaricarpa A. Nelson var. divaricarpa 2n = 13+2B, BC, Taylor et al. 3369 (DAO), Mulligan (1964). 2n = 14, MB, Live & Love 6026 (WIN), Love & Léve (1982) as Boechera divaricarpa. SK, Taylor & Sherk 4708 (DAO), Taylor & Brockman (1966). AB, Mosquin et al. 4703 & 4751 (DAO), Mosquin in Mulligan (1995). BC, Taylor & Ferguson 3054 (DAO), Mulligan (1964). 2n = 21, AB, Mosquin & Benn 5183 (DAO), Mosquin in Mulligan (1995). BC, Calder & Gillett 26501 & 27052 (DAO), Mulligan (1964). 2n = 22, PQ, Gervais 95-33 (QFA), Gervais et al. (1999) as Arabis holboellii. Arabis drummondii A. Gray 2n =14, YT, Calder & Kukkonen 28255 (DAO), Mulligan (1964). MB, Love & Love 6583 (WIN), Love & Love (1982) as Boechera drummondii. AB, Mosquin et al. 4706 & 4737 (DAO), Mosquin in Mulligan (1995). BC, Calder & MacKay 32750 (DAO), Mulligan (1964). 2n = 20, BC, Calder 28394 (DAO), Mulligan (1964). 2n = 21, BC, Mosquin et al. 4738 & 4755 (DAO), Mosquin in Mulligan (1995). Arabis eschscholtziana (Hopkins) Rollins 2n = 64, BC, Calder et al. 34754 (DAO), Taylor & Mulligan (1968) as Arabis hirsuta subsp. eschscholtziana. Arabis exilis A. Nelson 2n = 14, BC, Calder et al. 16531 & 16883 (DAO) and Calder & Spicer 33058 & 33427 (DAO), all in Mulligan (1964) as Arabis pendulocarpa; Gillett & Mitchell 3899 (DAO), Bécher (1969) as Arabis pendulocarpa; Taylor 6223 (UBC), Taylor & Taylor (1977), as Arabis holboellii var. pendulocarpa. 2002 MULLIGAN: CHROMOSOME NUMBERS OF MUSTARDS 613 Arabis glabra (L.) Bernh. (= Turritis glabra L.) 2n = 12, PQ, Gillett 10569 (DAO), Mosquin in Mulligan (1995). ON, Mulligan 2836 (DAO), Mulligan (1964) as Turritis glabra. AB, Mosquin & Mosquin 4687 (DAO), Mosquin in Mulligan (1995). BC, Calder & Taylor 36526 (DAO), Taylor & Mulligan (1968). Arabis hirsuta var. pycnocarpa (Hopkins) Rollins 2n = 32, NI-M, Cody & Spicer 11877 & 12249 (DAO), Mulligan (1964) as Arabis hirsuta. MB, Taylor & Sherk 4784 (DAO), Taylor & Brockman (1966) as Arabis hirsuta. AB, Mosquin et al. 4688, 4701 & 5184 (DAO), Mosquin in Mulligan (1995). BC, Mosquin & Mulligan 4933 (DAO), Mosquin in Mulligan (1995). Arabis holboellii var. consanguinea (Greene) G. A. Mulligan 2n = 21, AB, Mosquin et al. 4696, 4705 & 5212 (DAO) and Mosquin 5201 (DAO), Mosquin in Mulligan (1995). Arabis holboellii var. retrofracta (Graham) Rydb. 2n = 13+2B, MB, Boivin et al. 13209 (DAO), Mulligan (1964) as Arabis holboellii. 2n =14, MB, Gillett & Taylor 11693 (DAO), Mulligan (1964) as Arabis holboellii; Love & Love 6426 (WIN), Love & Love (1982) as Boechera collinsia; Mosquin & Mosquin 4911 (DAO), Mosquin in Mulligan (1995). SK, Boivin & Mosquin 10797 (DAO), Mulligan (1964) as Arabis holboellii; Mosquin & Mosquin 4914 & 4915 (DAO), Mosquin in Mulligan (1995). AB, Boivin & Perron 12626 (DAO), Mulligan (1964) as Arabis holboellii; Mosquin et al. 4454, 4460, 4683 & 4686 (DAO) and Calder & Gillett 26537 (DAO), all Mosquin in Mulligan (1995); Packer 6096 (ALTA), Packer (1964). BC, Calder et al. 26760, 26997 & 33417 (DAO), Mulligan (1964) as Arabis holboellii; Mosquin et al. 4454, 4460, 4929, 4935 & 4938 (DAO), Mosquin in Mulligan (1995). 2n = 14+1B, BC, Calder & Spicer 33794 (DAO), Mulligan (1964) as Arabis holboellii. 2n = 15, AB, Mosquin et al. 4735 (DAO), Mosquin in Mulligan (1995). BC, Mosquin & Mulligan 4932 (DAO), Mosquin in Mulligan (1995). 2n = 20+2B, SK, Mulligan 1049 (DAO), Mulligan (1964) as Arabis holboellii. 2n = ca.21, AB, Mosquin & Mulligan 4922 (DAO), Mosquin in Mulligan (1995). BC, Mosquin & Mosquin 4452 (DAO), Mosquin in Mulligan (1995). Arabis holboellii var. secunda (Howell) Jepson 2n = 21, BC, Calder & Spicer 33722A (DAO), Mulligan (1964) as Arabis holboellii. Arabis kamtschatica (Fisch. ex DC.) Ledeb. (= Arabidopsis lyrata subsp. kamchatica (Fisch. ex DC.) O’Kane & Al-Shehbaz) 2n = 32, AK, Drury 3380 (GH), Gaiser in Drury & Rollins (1952); Dawe (ALA), Dawe & Murray (1979) give the chromosome number of 2n = 16 for Arabis lyrata subsp. kamchatica but the voucher specimen in ALA clearly states that the number counted was 2n = 32. YT, R.T. Porsild 1275 (CAN), Mulligan & Porsild (1969a) as Arabis lyrata subsp. kamchatica; Porsild 1739 & 1788 (CAN), Mulligan & Porsild (1970) as Arabis lyrata subsp. kamchatica; Calder & Kukkonen 27785 (DAO), Mulligan (1964) as Arabis lyrata. BC, Calder & MacKay 30637 & 31670A (DAO) and Calder et al. 18457 (DAO), Mulligan (1964) as Arabis lyrata; Calder & Taylor 36389 (DAO), Taylor & Mulligan (1968) as Arabis lyrata subsp. kamchatica. Arabis lemmonii S. Watson 2n = 14, AB, Calder & MacKay 32722A (DAO), Mulligan (1964). BC, Taylor et al. 3250 (DAO), Mulligan (1964). Arabis lyallii S. Watson 2n = 21, AB, Calder & MacKay 32671 (DAO), Mulligan (1964). BC, Taylor & Ferguson 3815 (DAQ), Mulligan (1964). Arabis lyrata L. (= Arabidopsis lyrata (L.) O’Kane & Al-Shehbaz subsp. /yrata) 2n = 16, ON, Senn & Lindsay 5399 (DAO), Bocher (1969). SK, Boivin & Mosquin 10769 (DAQ), Mulligan (1964). AB, Packer 80-29 (ALTA), Packer & Witkus (1982) as Arabis lyrata subsp. kamchatica. Arabis microphylla Nutt. ex Torr. & Gray 2n = 14, BC, Taylor & Ferguson 2641 (DAO), Mulligan (1964). Arabis petraea (L.) Lam. (= Arabis media N. Busch) 2n = 16, AK, Hogdon 8664 (GH), Rollins (1966) as Arabis lyrata var. kamchatica; Packer (ALTA), Johnson & Packer (1968) as Arabis lyrata subsp. kamchatica. Arabis pinetorum Tidestr. 2n = 13+2B, MB, Boivin et al. 13209 (DAO), Mulligan (1964) as Arabis holboellii. 2n = 14, AK, Batten & Dawe 78-192A (ALA), Dawe & Murray (1979) as Arabis holboellii. 2n = 32, BC, Taylor 6231 (UBC), Taylor & Taylor (1977) as Arabis sparsiflora var. columbiana. 614 THE CANADIAN FIELD-NATURALIST Vol. 116 Athysanus pusillus (Hook.) Greene 2n = 26, BC, Calder & MacKay 30873 (DAO), Mulligan (unpublished). Barbarea orthoceras Ledeb. 2n = 16, AK, Hatch 79 (ALA), Dawe & Murray (1979). YT, Porsild 546 (CAN), Mulligan & Porsild (1969a). BC, Calder & MacKay 31716 (DAO), Mulligan (1964); Calder et al. 34756 & 34847 (DAO), Taylor & Mulligan (1968). Barbarea vulgaris (L.) W. T. Aiton 2n = 16, PQ, Gervais 71-030 (QUE), Gervais (1979). ON, Mulligan 2286 (DAO), Mulligan (1959); Grainger & MacLeod 4032, 4035, 4040, 4061, 4067, 4081, 4084, 4125, 4136 & 4172 (DAO), Mulligan & Grainger 4007 (DAO), Mulligan et al. 4018 & 4051 (DAO) and MacLeod et al..4025 (DAO), all in Mulligan (1984). Berteroa incana (L.) DC. 2n = 16, PQ, Cayouette 1720 (QUE), Gervais (1979). ON, Mulligan 2125 (DAO), Mulligan (1957). Brassica juncea (L.) Czern. 2n = 36, AB, Mulligan 1483 (DAO), Mulligan (1959). Brassica rapa L. (= Brassica campestris L.) 2n = 20, NB, Mulligan 1482 (DAO), Mulligan (1959) as Brassica campestris. PQ, Moreau (QUE), Gervais (1979). BC, Calder & Taylor 36051 (DAO), Taylor & Mulligan (1968) as Brassica campestris. Braya glabella Richardson (= Braya purpurascens (R. Br.) Bunge) 2n = 56, AK, Packer 2098 (ALTA), Johnson & Packer (1968) as Braya purpurascens; Dawe & Lipkin 848 (ALA), Dawe & Murray (1979) as Braya purpurascens; Kelso 83-209a (ALA), Murray & Kelso (1997); Murray 3518 (DAO), Dawe (unpublished). N-F, Savile 4764A & 4707 (DAO), Mulligan (1965) as Braya purpurascens. BC, Calder & Kukkonen 27446 (DAO), Mulligan (1965) as Braya purpurascens. Braya humilis (C. A. Mey.) B. L. Rob. (= Neotorularia humilis (Ledeb.) Hedge) 2n = 28, NT-M, Cody & Spicer 12128 (DAO), Mulligan (1965); Cody & Scotter 19268 (DAO), Mulligan et al. (1972). AB, Mosquin & Mosquin 4691(DAO), Mulligan (unpublished). BC, Taylor & Ferguson Z2427 (DAO), Mulligan (1965). 2n = 42, YT, Porsild 1701 (CAN), Mulligan & Porsild (1970). NT-M, Cody & Matte 9039 & 9334 (DAO), Cody & Gutteridge 7339 (DAO), Cody 13166 (DAO), Cody & Spicer 12259 (DAO) and Cody & Kehoe 12692, 12723 & 12732 (DAO), all in Mulligan (1965). N-F, Savile 4669 & 4763 (DAO), Mulligan (1965). AB, Calder & Spicer 33838 (DAO), Mulligan (1965). BC, Calder & Savile 11979 (DAO) and Calder & Kukkonen 27375 (DAO), both in Mulligan (1965). 2n = 56, NT-M, Cody & Gutteridge 7774 (DAO), Mulligan (1965). BC, Calder 28360 (DAO) and Taylor & Ferguson 3899 (DAO), both in Mulligan (1965). Braya thorild-wulfii Ostenf. 2n = 28, N-F, Savile 4456, 4764B & 4740 (DAO) and Beschel 11070 (DAO), all in Mulligan (1965). Cakile edentula (Bigelow) Hook. var. edentula 2n = 18, PQ, Bassett & Crompton 4266 (DAO), Mulligan (1964); Cody & Kemp 18194 (DAO), Mulligan (unpublished). BC, Calder & MacKay 31917 & 31959 (DAO), Mulligan (1964); Calder & Taylor 35336 (DAO),Taylor & Mulligan (1968). Cakile edentula var. lacustris Fernald 2n = 18, ON, Gillett 10723 (DAO), Mulligan (1964) as Cakile edentula. Cakile maritima Scop. 2n = 18, BC, Calder & Taylor 35430 (DAO), Taylor & Mulligan (1968). Camelina microcarpa Andrz. ex DC. 2n = 40, AB, Mulligan 2108 (DAO), Mulligan (1957). SK, MacLeod 188 (DAO), Mulligan (1984). Camelina sativa (L.) Crantz. 2n = 40, AB, Groh 2805 (DAO) and Frankton 1485 (DAO), both R. J. Moore (unpublished). BC, Groh 2997 (DAO) and Frankton 1486 (DAO), both Moore (unpublished). Capsella bursa-pastoris (L.) Medik. 2n = 32, ON, Mulligan 1493 & 1523 (DAO), Mulligan (1957); Grainger & MacLeod 4002, 4037, 4045, 4062, 4090, 4107, 4120, 4124, 4182, 4184 & 4187 (DAO) and Mulligan et al. 4017 (DAO), all in Mulligan (1984). SK, MacLeod 44 (DAO), Mulligan (1984). BC, Calder & Taylor 35168 (DAO), Taylor & Mulligan (1968). Cardamine angulata Hook. 2n = 40, BC, Calder et al. 34681 & 35030 (DAO) and Calder & Taylor 35497 (DAO), all Mulligan (1965). 2002 MULLIGAN: CHROMOSOME NUMBERS OF MUSTARDS 615 Cardamine bellidifolia L. 2n = 16, AK, Packer 1832 (ALTA), Johnson & Packer (1968); McPherson & Galeski 72-9 (ALTA), Packer & McPherson (1974); Murray & Johnson 6310 (ALA), Dawe & Murray (1979). YT, Porsild 1264 (CAN), Mulligan & Porsild (1969a). N-F, Savile 4496 (DAO), Mulligan (1965). PQ, Trahan 92-43 (QFA), Gervais et al. (1997). AB, Packer, May 1959 (ALTA), Packer (1964). BC, Calder & MacKay 32513 (DAO) and Calder & Savile 36447 (DAO), both in Mulligan (1965). Cardamine cocatenata (Michx.) O. Schwartz 2n = ca. 240, ON, Montgomery 195 (OAC), Montgomery (1955) under Dentaria lanciniata. Cardamine digitata Richardson 2n = 28, AK, Thompson (GH), Rollins (1966); Packer 1838 (ALTA), Johnson & Packer (1968). Cardamine diphylla (Michx.) A. W. Wood 2n = 96, ON, Montgomery 194 (OAC), Montgomery (1955) as Dentaria diphylla. Cardamine hirsuta L. 2n = 16, BC, Calder et al. 16209 & 16457 (DAO), Mulligan (1965). Cardamine microphylla Adams 2n = 28, AK, Kelso et al. 63 (ALA), Murray & Kelso (1997) as Cardamine microphylla subsp. blaisdellii. 2n = 64, NT-M, Calder 34009 (DAO), Mulligan (1965). Cardamine occidentalis (S. Watson) Howell 2n = 64, BC, Calder & Taylor 35271 & 37465 (DAO), Mulligan (1965). Cardamine oligosperma Nutt. ex Torr. & Gray 2n = 16, BC, Lindsay & Woodbury 647 (DAO) and Calder et al. 20895, 21073 & 22219A (DAO), all in Mulligan (1965). Cardamine parviflora var. arenicola (Britton) O. E. Schulz 2n = 16, ON, Dore 19111 (DAO), Mulligan (1965) as Cardamine parviflora. Cardamine pensylvanica Muhl ex Willd. 2n = 32, YT, Calder 26547 (DAO), Mulligan (1965). NB, Cunningham (DAO), Mulligan (1965). PQ, Dore 18633 (DAO), Mulligan (1965); Gagnon 96-183 (QFA), Gervais et al. (1999) as Capsella bursa- pastoris. BC, Calder & Taylor 37514 (DAO) and Calder et al. 19964 & 22580 (DAO), all in Mulligan (1965). 2n = 64, ON, Dore 18681 (DAO), Mulligan (1965). Cardamine pratensis L. 2n = 80 to 100, AK, McPherson & Galeski 72-258 (ALTA), Packer & McPherson (1974). Cardamine purpurea Cham. & Schlecht. 2n = ca.80, AK, Packer 2555 (ALTA), Johnson & Packer (1968). Cardamine umbellata Greene 2n = 48, BC, Taylor & Calder 3751 (DAO), Calder & MacKay 31402 (DAO), Calder & Taylor 35966 & 36297 (DAO) and Calder et al. 22226, 34730 & 34755 (DAO), all in Mulligan (1965). Cardaria chalepensis (L.) Hand.-Mazz. 2n = 80, SK, Mulligan 2408, 2409 & 2410 (DAO), Mulligan & Frankton (1962). AB, Mulligan 2432, 2433, 2441, 2442, 2443, 2444, 2446, 2447 & 2576 (DAO), Mulligan & Frankton (1962). BC, Mulligan 2402 (DAO), Mulligan & Frankton (1962). Cardaria draba (L.) Desv. 2n = 64, AB, Mulligan 2435, 2436 & 2437 (DAO), Mulligan & Frankton (1962). Cardaria pubescens (C. A. Mey.) Jarmol. 2n = 16, SK, Mulligan 2403, 2429, 2430 & 2431 (DAO), Mulligan & Frankton (1962). AB, Mulligan 2440 (DAO), Mulligan & Frankton (1962). Cochlearia groenlandica L. 2n = 14, AK, Baldwin 1894 (GH), Rollins (1966); McPherson & Galeski 72-6 (ALTA), Packer & McPherson (1974) as Cochlearia officinalis; Kelso et al. 66 (ALA), Murray & Kelso (1997); Packer (ALTA), Johnson & Packer (1968). NT-M, Parmelee 2967 & 3203 (DAO), Mulligan (unpublished). N-F, Hedberg 3246 (UPS), Hedberg (1967); Mosquin & Martin 6444 (DAO), Mosquin & Hayley (1966) as Cochlearia officinalis. BC, Calder & Taylor 34727 & 35141 (DAO) and Calder et al. 22231 (DAQ), all in Taylor & Mulligan (1968) as Cochlearia officinalis; Emrich 84 (DAQ), Mulligan (unpublished). Descurainia incana (Bernh. ex Fisch. & C. A. Mey.) Dorn (= Descurainia richardsonii (Sweet) O. E. Schulz) 2n = 14, YT, Gillett 4497 (DAO), Mulligan (1961a) as Descurainia ric thardsonii. PQ, Anderson 1220 (DAO), Mulligan (1961a) as Descurainia richardsonii. MB, A. Live 5216 (WIN), Live & Live (1982) as Descurainia richardsonii. 616 THE CANADIAN FIELD-NATURALIST Vols; Descurainia incisa (Engelm. ex Gray) Britton var. incisa - 2n = 14, YT, Packer 1500 (ALTA), Packer (1964) as Descurainia sophioides. Descurainia pinnata var. brachycarpa (Richardson) Fernald 2n = 28, MB, Live & Love 5853 (WIN), Live & Love (1982) as Descuria pinnata. AB, Mulligan 2126 (DAO), Mulligan (1961a). Descurainia sophia (L.) Webb. 2n = 28, SK, MacLeod 212 (DAO), Mulligan (1984). AB, Mulligan 2127 (DAO), Mulligan (1961a). BC, Calder & Taylor 36036 (DAO), Taylor & Mulligan (1968). Descurainia sophioides (Fisch. ex Hook.) O. E. Schulz 2n = 14, YT, Calder 4216 (DAO), Mulligan (1961a). N-F, Kuc, July 25, 1968 (CAN), Mulligan & Porsild (1970). Diplotaxis tenuifolia (L.) DC. 2n = 22, ON, Mulligan 2273 (DAO), Mulligan (1959). Draba albertina Greene 2n = 24, NT-M, Cody 16443 (DAO), Mulligan (1975). AB, Mulligan et al. 3350 (DAO), Mulligan (1975). BC, Mulligan & Crompton 3611, 3612 & 3616 (DAO), in Mulligan (1975). Draba alpina L. 2n = 80, AK, Packer et al. 72-16a (ALTA, photo DAO), Packer & McPherson (1974). N-F, Mulligan 3560, 3562 & 3563 (DAO), Mulligan (unpublished); Mulligan 3535 & 3542 (DAO) and Maycock & Op de Beeck 10493 (DAO), all in Mulligan (unpublished). Draba arabisans Michx. ; 2n = ca.94, PQ, Gervais 95-184 (QFA), Gervais et al. (1999). 2n = 96, PQ, Mulligan 2917 (DAO), Mulligan & Beales 3147, 3196 & 3202 (DAO) and Cody & Kemp 18295 (DAO), all in Mulligan (1970a). Draba aurea M. Vahl 2n = ca.72, PQ, Gervais 95-183B (QFA) and Gervais 95-188C (QFA), both in Gervais et al. (1997). 2n = 74, YT, Mulligan 3564 (DAO), Mulligan (unpublished). PQ, Dutilly & Lepage 39413 (DAO), Mulligan (1966); Mulligan & Beales 3201 (DAO), Mulligan (unpublished). BC, Mulligan & Crompton 3239 & 3251 (DAO), Mulligan (unpublished). Draba borealis DC. 2n = 80, YT, Porsild 1276 (CAN) and Mulligan 3204 (DAO), Mulligan (1970a). AB, Calder 23936 (DAO), Mulligan (1966) as Draba McCallae; Mulligan & Mulligan 3379 & 3501 (DAO), Mulligan (1970a). 2n = 82, AB, Calder 37270 (DAO), Mulligan (1966) as Draba McCallae; Mulligan & Mulligan 3384 (DAO), Mulligan (unpublished). Draba cana Rydb. 2n = 32, AK, Murray & Murray 2759 (DAO), Mulligan (1971a). YT, Calder & Kukkonen 28213 (DAO), Mulligan (1971a). NT-M, Kvale & Haggard 52 (DAO) and Calder 17244 (DAO), both in Mulligan (1971a). PQ, Cody & Mulligan 16325 (DAO) and Frankton & Frankton 2044 (DAO), both in Mulligan (1971a). AB, Mulligan 2918 (DAO), Mulligan & Crompton 3228 & 3261 (DAO) and Mulligan & Mulligan 3319A, 3470, 3486, 3499 & 3503A (DAO), all in Mulligan (1971a). BC, Calder & Kukkonen 26906 (DAO), Mulligan (197 1a). Draba cinerea Adams 2n = 48, AK, Kelso 84-381 (ALA), Murray & Kelso (1997). NT-M, Cody & Kehoe 12676A (DAO), Mulligan (1966) as Draba lanceolata; Scotter 7189 (DAO) and Cody 16058 (DAO), both in Mulligan (1971a). N-F, Savile 4667 (DAO), Mulligan (197 1a). Draba corymbosa R. Br. ex DC. 2n = ca.110 to 130, Mosquin & Martin 6472 (DAO), Mosquin & Hayley (1966) as Draba bellii. 2n = 144, NT-M, Parmelee 3205 (DAO) in Mulligan (1976). N-F, Parmelee & Seaborn 3919A (DAO) and Mulligan 3261(DAO), both in Mulligan (1976). Draba crassifolia Graham 2n = 40, YT, Porsild 1952 (CAN), Mulligan 3510 (DAO) and Scotter 21025 (DAO), all in Mulligan (1975). AB, Calder & MacKay 32714 (DAO), Mulligan (1966); Mulligan & Mulligan 3397, 3444 & 3492 (DAO), Mulligan (1975). BC, Taylor & Ferguson 3760 (DAO), Mulligan (1966); Taylor et al. 5603 (DAO), Mulligan (1975). Draba densifolia Nutt. 2n = 36, AB, Mulligan & Crompton 3245 (DAO), Mulligan (1976).Draba fladnizensis Wulf. 2n = 16, YT, Scotter 20707, 20862 & 20875 (DAO), Mulligan (1974a). Nm MULLIGAN: CHROMOSOME NUMBERS OF MUSTARDS 617 tba glabella Pursh var. glabella 2a = 64, AK, Kelso 83-272 (ALA), Murray & Kelso (1997). YT, Porsild 227, 228 & 1677 (CAN), Mulligan (1970a). NT-M, Cody 15818 & 17627 (DAO) and Scotter 7181 (DAO), all in Mulligan (1970a); Kuc 410 (DAO) and Mulligan 3561 (DAO), Mulligan (unpublished). N-K, Maycock & Op de Beeck 9649, 9664, 10046, 10624, 10956 & 11057 (DAO) and Mulligan 3512, 3514, 3515, 3516, 35 16a, 3519, 3520 (unusual, with creamy-yellow flowers), 3521, 3540, 3543 & 3547 (DAO), all in Mulligan (1970a). N-F, Mason 91 & 112 (DAO), Mulligan (1971a) as Draba cinerea. PQ, Mulligan & Beales 3144, 3145, 3148, 3150, 3166, 3179, 3182, 3183, 3184, 3190, 3195, 3199 & 3211 (DAO) and Lemieux et al. 3889 (DAO), all in Mulligan (1970a). 2n = 80, AK, Packer 1780b, 1926 & 1956 (ALTA), Johnson & Packer (1968) as Draba hirta. N-K, Maycock & Op de Beeck 9937 (DAO) and Mulligan 3533 (DAO), Mulligan (1970a). Draba glabella var. pycnosperma (Fernald & Knowiton) G. A. Mulligan | 2n = 64, PQ, Sherk & Cing-Mars 480 (DAO), Mulligan (1970a) as Draba glabella. Draba grandis Langsd. (= Draba hyperborea (L) Desv. and Nesodraba grandis (Langsd.) E. L. Greene) 2n = 36, BC, Calder & MacKay 31393 (DAO), Mulligan (1966) as Draba hyperborea. 2n = 38, BC, Calder & Taylor 36570 (DAO), Mulligan (1966) as Draba hyperborea. Draba incana L. 2n = 32, PQ, Sherk & Cing-Mars 483 (DAO), Mulligan (1970a). Draba incerta Payson 2n = 112, YT, Murray 1185 (ALA,DAO), Mulligan (1972). AB, Calder & MacKay 32710 (DAO), Mulligan (1966); Mulligan & Crompton 3235 & 3244 (DAO), Calder 37194, 37299 & 37307 (DAO) and Mulligan & Mulligan 3353, 3473, 3481, 3485 & 3495 (DAO), all in Mulligan (1972). BC, Taylor et al. 3324 (DAO), Mulligan (1966). Draba juvenilis Kom. (= Draba longipes Raup) 2n = 64, YT, Porsild 1277 & 1278 (CAN) and Mulligan 3262 & 3263 (DAO), Mulligan (1970a) as Draba longipes. NT-M, Cody 16589 & 16616 (DAO), Mulligan (1970a) as Draba longipes. Draba kananaskis G. A. Mulligan 2n = 64, AB, Mulligan & Mulligan 3477 & 3548 (DAO), Mulligan (1970b). Draba lonchocarpa Rydb. var. lonchocarpa 2n = 16, YT, Porsild 1698 (CAN), Mulligan (1974a). AB, Calder & MacKay 32710A (DAO), Mulligan & Mulligan 3474 & 3488 (DAO) and Mulligan & Crompton 3219 (DAO), all in Mulligan (1974a). BC, Calder & Kukkonen 27216 (DAO), Calder & MacKay 32771 (DAO) and Mulligan & Mulligan 3459 (DAO), all in Mulligan (1974a). Draba lonchocarpa var. vestita O. E. Schulz 2n = 16, BC, Calder & Taylor 36382 (DAO), Mulligan (1974a). Draba macounii O. E. Schulz 2n = 64, BC, Mulligan & Mulligan 3450 (DAO) and Calder & Kukkonen 27184 (DAO), Mulligan (1976). Draba murrayi G. A. Mulligan 2n = 48, AK, Batten & Dawe 78-291 (ALA, photo DAO), Dawe & Murray (1981b). Draba nemorosa L. 2n = 16, ON, Frankton 1915 (DAO) and Cody 18059 (DAO), both in Mulligan (1975). MB, Boivin & Love 5404 (WIN), Love & Love (1982). SK, Mulligan 2835 (DAO) and Sherk & Taylor 134 (DAQ), both in Mulligan (1975). AB, Packer, June 2, 1962 (ALTA), Packer (1964); Mosquin & Mosquin 4685 (DAO), Mulligan (1966); Mulligan & Crompton 3223 (DAO), Mulligan (1975). Draba nivalis Liljebl. 2n = 16, AK, Packer, 12 Aug., 1962 (ALTA), Johnson & Packer (1968); Dawe 79-52 (ALA), Dawe & Murray (1980); Batten et al. 78-67 (ALA), Dawe & Murray (1981b) as Draba lactea. YT, Porsild 464 (CAN), Mulligan & Porsild (1969a). NT-M, Cody 17628 (DAO) and Scotter 6462a (DAQ), both in Mulligan (1974a). N-F, Parmelee & Seaborne 4027 & 4177 (DAO), Mulligan (1974a). N-K, Whitton, Aug. 30, 1967 (DAO) and Maycock & Op de Beeck 9832, 10805, 11014 & 11073 (DAO), all in Mulligan (1974a). BC, Taylor 5713 (DAO), Mulligan (1974a). Draba norvegica R. Br. ex DC. 2n = 48, NT-M, Cody 16016 (DAO), Mulligan & Cody (1968). N-K, Maycock & Op de Beeck 10165, 10405, 10500B, 10535 & 10934 (DAO), all in Mulligan ( 1970a). PQ, Trahan 92-92A (QFA), Gervais et al. (1997) under Draba clivicola. Draba oblongata R. Br. ex DC. (= Draba groenlandica E. Ekman) 2n = 64, N-F, Parmelee 2089 (DAO), Mulligan (197 1a) as Draba groenlandica. 618 THE CANADIAN FIELD-NATURALIST Vol. 116 Draba ogilviensis Hultén 2n = 16, YT, Porsild 141 (CAN), Mulligan & Porsild (1969b) as Draba sibirica. Draba oligosperma Hook. 2n = 64, YT, Porsild 1336 (CAN), Mulligan (1972). AB, Mulligan & Crompton 3217, 3221, 3242, 3243, 3246 & 3255 (DAO) and Mulligan & Mulligan 3471(DAO), all in Mulligan (1972). Draba pauciflora R. Br. (= Draba adamsii Ledeb.) 2n = 32, AK, Packer et al. 72-80 (ALTA), Packer & McPherson (1974) as Draba adamsii; Packer et al. 72-16b & 72-212 (ALTA, photos DAO), Mulligan (1974b) as Draba adamsii. Draba paysonii J. F. Macbr. 2n = 42, AB, Mulligan & Crompton 3230 & 3249 (DAO) and Mulligan & Mulligan 3333 (DAO), Mulligan (1971b). Draba porsildii G. A. Mulligan 2n = 32, YT, Scotter, Aug. 4,1972 (DAO), Mulligan (1974b). AB, Mulligan & Crompton 3220 (DAO), Calder & Kukkonen 27184A (DAO), Calder 37299a (DAO) and Mulligan & Mulligan 3472 & 3491 (DAO), all in Mulligan (1974a). Draba praealta Greene 2n = 56, AB, Calder 37269 (DAO), Mulligan (1966). BC, Taylor & Ferguson 3187 (DAO), Mulligan (1966). Draba reptans Lam. 2n = 30, SK, Boivin 13494 (DAO), Mulligan (1966). Draba ruaxes Payson & H. St. John 2n =ca.72, YT, Murray & Murray 1343 (ALA,DAO), Mulligan (1971b). Draba scotteri G. A. Mulligan 2n = 96, YT, Scotter 21041A (DAO), Mulligan (1979). Draba stenoloba Ledeb. 2n = 40, AB, Scotter 13993 (DAO), Mulligan 3558 (DAO) and Mulligan & Mulligan 3498 (DAO), all in Mulligan (1975). Draba subcapitata Simmons 2n = 16, N-F, Mason 69 (DAO) and Mulligan 3511 (DAO), Mulligan (1974a). Draba ventosa A. Gray “ 2n = 36, AB, Mulligan & Mulligan 3489 (DAO), Mulligan (1971b). BC, Mulligan & Mulligan 3466 (DAO), Mulligan (1971b). Draba verna L. 2n = 39, BC, Calder & Spicer 33089 (DAO), Mulligan (1966). Draba wahlenbergii Hartm. (= Draba lactea of most authors) 2n = 48, AK, Packer et al. 72-1 (ALTA), Packer & McPherson (1974) as Draba lactea; Young 1380 (DAO) and Murray & Murray 2267 (ALA, DAO), both in Mulligan (1974a) as Draba lactea; the voucher specimens for the 2n = 16 count listed in Dawe & Murray (1981b), Batten et al. 78-67 (ALA), are all specimens of Draba nivalis. N-F, Parmelee & Seaborne 3760 & 3922 (DAO), Mulligan (1974a) for Draba lactea. N-K, Maycock & Op de Beeck 10335, 10482 &10751 (DAO), Mulligan 3526, 3544 & 3559 (DAO) and Morisette 70/156 (DAO), all in Mulligan (1974a) as Draba lactea. Erucastrum gallicum (Willd.) O. E. Schulz 2n = 30, MB, Mulligan 2128 (DAO), Mulligan (1957). SK, MacLeod 168 (DAO), Mulligan (1984). Erysimum angustatum Rydb. 2n = 36, AK, Batten & Dawe 78-220 (ALA), Dawe & Murray (1981b). Erysimum arenicola §. Watson 2n = 36, BC, Calder & MacKay 32024 & 32184 (DAO), Mulligan (1966) as Erysimum capitatum. Erysimum cheiranthoides L. 2n = 16, PQ, Gervais 71-341 (QUE), Gervais (1981). ON, Mulligan 1444 (DAO), Mulligan (1957); Mulligan & Grainger 4005 (DAO), Mulligan (1984). BC, Calder & Taylor 36026 (DAO), Taylor & Mulligan (1968). Erysimum coarctatum Fernald 2n = 54, YT, Martin 24 & 40 (DAO), Calder & Kukkonen 27983 & 28014 (DAO), Calder 28352 (DAO) and Cody 12564 (DAO), all in Mulligan (1966) as Erysimum inconspicuum. Erysimum hieraciifolium L. 2n = 48, ON, Mulligan & Rae 2137 (DAO), Mulligan (1959); Mulligan & Mulligan 3103 & 3106, meiosis irregular and 4, 5, or 6 microspores (DAO), Mulligan & Frankton (1967). Erysimum inconspicuum (S. Watson) MacMill. 2n = 81, ON, Dore & Gillett 17515 (DAO) and Garton 6807 (DAO), both in Mulligan (1966). MB, Bassett 3580 (DAO), Mulligan (1966). AB, Jenkins 7954 (DAO), Mulligan (1966). BC, Mosquin 2678 & 2681 (DAO), Mulligan (1966). 2002 MULLIGAN: CHROMOSOME NUMBERS OF MUSTARDS 619 Erysimum pallasii (Pursh) Fernald 2n = 24, YT, Calder 34130 & 34295 (DAO), Mulligan (1966); Packer 1385 (ALTA), Packer (1964). N-F, Mosquin & Martin 6405 (DAO), Mosquin & Hayley (1966). 2n = 28, YT, Porsild 55 (CAN), Mulligan & Porsild (1969a). 2n = 36, AK, Thompson (DS), Rollins (1966). 2n = 42, AK, Hulten (S), Hedberg (1967). Eutrema edwardsii R. Br. 2n = 28, AK, Packer (ALTA), Johnson & Packer (1968); Packer et al. 72-66 (ALTA), Packer & McPherson (1974). N-F, Savile 4724 (DAO), Mulligan (1964); Hedberg 3305 (UPS), Hedberg (1967). 2n = 42, N-F, Parmelee 2200a (DAO), Mulligan (1964). Halimolobos mollis (Hook.) Rollins 2n = 16, YT, Calder & Gillett 24966 (DAO), Mulligan (1964). 2n = 20, AK, Batten & Dawe 78-384 (ALA), Dawe & Murray (198 1a). Hesperis matronalis L. 2n = 24, ON, Cody 21126 (DAO), Mulligan (1984). Lepidium bourgeauanum Thell. 2n = 32, NT-M, Mulligan 2418 & 2419 (DAO), Mulligan (1961a,b). PQ, Mulligan 2423 (DAO), Mulligan (1961a,b). Lepidium campestre (L.) R. Br. 2n = 16, ON, Mulligan 1499 (DAO), Mulligan (1957). SK, Mulligan 2219 (DAO), Mulligan (1957). BC, Calder & Taylor 36012 (DAO), Taylor & Mulligan (1968). Lepidium densiflorum Schrad. var. densiflorum 2n = 32, PQ, Gervais 77-101 (QUE), Gervais (1979). ON, Mulligan 1528 (DAO), Mulligan (1957); Mulligan 2413 (DAO), Mulligan (1961a,b). SK, MacLeod 233 (DAO), Mulligan (1984). Lepidium densiflorum var. elongatum (Rydb.) Thell. 2n = 32, BC, Mulligan 2422 (DAO), Mulligan (1961a,b). Lepidium densiflorum var. macrocarpum G. A. Mulligan 2n = 32, BC, Mulligan 2416 (DAO), Mulligan (1961a,b). Lepidium latifolium L. 2n = 24, AB, Mulligan 2147 (DAO), Mulligan (1957). Lepidium perfoliatum L. 2n = 16, AB, Mulligan 1577 (DAO), Mulligan (1957). BC, Stein 4302 (UBC), Taylor & Taylor (1977). Lepidium ramosissimum A. Nelson 2n = 64, NT-M, Mulligan 2417 (DAO), Mulligan (1961a,b). SK, MacLeod 27 (DAO), Mulligan (1984). AB, Mulligan 2129 (DAO), Mulligan (1957); Mulligan 2424 (DAO), Mulligan (1961a,b). Lepidium virginicum L. 2n = 32, ON, Mulligan 2420 (DAO), Mulligan (1961a,b). BC, Mulligan 2421 (DAO), Mulligan (1961a,b). Lesquerella arctica (Wormsk. ex Hornem.) S. Watson 2n = 60, AK, Murray 6727 (ALA), Dawe & Murray (1981a). N-F, Savile 4459 (DAO) and Bruggeman 4459 (DAO), both Mulligan in Rollins & Shaw (1973). Lesquerella arenosa (Richardson) Rydb. 2n = 10, SK, Mulligan & Mosquin 4913 (DAO), Mulligan in Rollins & Shaw (1973). AB, Mulligan & Mosquin 2670 (DAO), Mulligan in Rollins & Shaw (1973). Lesquerella calderi G. A. Mulligan & A. Porsild 2n = 20, YT, Porsild 1095 (CAN) and Mulligan 3212 & 3213 (DAO), Mulligan & Porsild (1969b). Lesquerella douglasii S. Watson 2n = 10, BC, Mulligan & Mosquin 2683 (DAO) and Calder & Spicer 32860 (DAO), Mulligan in Rollins & Shaw (1973). Nasturtium crystallinum (Rollins) G. A. Mulligan 2n = 32, NT-M, Cody 14789 (DAO), Mulligan & Cody (1995). Nasturtium microphyllum Boenn. ex Rchb. 2n = 64, PE, Erskine 2487 (DAO), Mulligan (1964). ON, Mulligan & Moore 2651 (DAQ), Mulligan (1964); Cody 21136 (DAO), Mulligan (1984). Neslia paniculata (L.) Desv. 2n = 14, ON, Mulligan 1485 (DAO), Mulligan (1957). BC, Calder & Taylor 36018 (DAQ), Taylor & Mulligan (1968). Parrya arctica R. Br. 2n = 14, N-F, Mosquin & Martin 6471 (DAO), Mosquin & Hayley (1966). 2n = ca.21, N-F, Mosquin & Martin 6401 (DAO), Mosquin & Hayley (1966). 620 THE CANADIAN FIELD-NATURALIST Vol. 116 Parrya nudicaulis (L.) Regel 2n = 14, N-F, Sgrensen (UPS), Hedberg (1964). 2n = 28, YT, Porsild 1683 (CAN), Mulligan & Porsild (1970).There is no voucher specimen for the count of 2n = 28 in Johnson & Packer (1968) according to J. G. Packer in personal correspondence, 13 June 2001. Physaria didymocarpa (Hook.) A. Gray 2n = 16, AB, Calder 23921 (DAO) and Mulligan & Mosquin 2669 & 2672 (DAO), all in Mulligan (1968). Raphanus raphanistrum L. 2n = 18, NB, Mulligan 2555 (DAO), Mulligan (1961a). Rorippa amphibia (L.) Besser 2n = 32, PQ, Gervais 96-17 (QFA), Gervais et al. (1999) as Rorippa sylvestris. Rorippa barbareifolia (DC.) Kitagawa 2n = 16, AK, Hatch 79 (ALA), Dawe & Murray (1979). YT, Porsild 547B & 1209 (CAN), Mulligan & Porsild (1968); Porsild 1722 (CAN), Mulligan & Porsild (1970). Rorippa curvipes Greene var. curvipes 2n = 16, BC, Calder & Taylor 35945 (DAO), Taylor & Mulligan (1968) as Rorippa islandica. Rorippa curvisiliqua (Hook.) Bessey ex Britton 2n = 16, BC, Molyneux 223 (DAO), Mulligan (1964). Rorippa palustris (L.) Besser 2n = 32, AK, Murray & Johnson 6309 (ALA), Dawe & Murray (1980). YT, Calder & Kukkonen 28028 & 28299 (DAO), Mulligan (1964) as Rorippa islandica. NS, Smith et al. 15009 (DAO), Mulligan (1964) as Rorippa islandica. PQ, Rolland-Germain 7012 (DAO), Mulligan (1964) as Rorippa islandica. MB, Bassett & Kemp 3536 (DAO), Mulligan (1964) as Rorippa islandica; Love & Léve 5486 & 5820 (WIN), Love & Love (1982). SK, Selleck 303 (DAO), Mulligan (1964) as Rorippa islandica; MacLeod 103 (DAO), Mulligan (1984). BC, Eastham 566 (DAO), Mulligan (1964) as Rorippa islandica. Rorippa sylvestris (L.) Besser 2n = 32, PQ, Mulligan 2527 (DAO), Mulligan & Munro (1984). ON, Mulligan 2537, 2538 & 2580 (DAO), Mulligan & Munro (1984). 2n = 40, ON, Mulligan 2533, 2534 & 2535 (DAO), Mulligan & Munro (1984). BC, Mulligan 2539 (DAO), Mulligan & Munro (1984). 2n = 48, NB, Mulligan, 2515, 2516, 2517 & 2518 (DAO), Mulligan & Munro (1984). PQ, Mulligan 2509, 2510, 2514, 2519, 2521, 2522, 2523, 2524, 2525 , 2528 & 2530 (DAO), Mulligan & Munro (1984), ON, Mulligan 2501, 2505, 2506, 2508, 2511, 2512, 2531, 2532 & 2536 (DAO), Mulligan & Munro (1984). AB, Mulligan 2270 (DAO), Mulligan & Munro (1984). BC, Mulligan 2268, 2278 & 2279 (DAO), Mulligan & Munro (1984). Schoenocrambe linifolia (Nutt.) Greene 2n = 14, BC, Taylor 6228 (UBC), Taylor & Taylor (1977). Sinapis arvensis L. 2n = 18, ON, Grainger & MacLeod 4177, 4228, 4324 & 4352 (DAO) and MacLeod 4834 (DAO), all in Mulligan (1984). MB, Warwick et al. 8154, 8221 & 8224 (DAO), Warwick et al. (1994). SK, MacLeod 18 & 99 (DAO), Mulligan (1984). AB, Warwick et al. 8225 & 8226 (DAO), Warwick et al. (1994). BC, Calder & Taylor 36014 (DAO), Taylor & Mulligan (1968). Sisymbrium altissimum L. 2n = 14, ON, Mulligan 1471 (DAO), Mulligan (1961a). SK, MacLeod 89 & 174 (DAO), Mulligan (1984). BC, Calder & Taylor 36043 (DAO), Taylor & Mulligan (1968). Sisymbrium loesellii L. 2n = 14, AB, Mulligan 2130 (DAO), Mulligan (1957). Sisymbrium officinale (L.) Scop. 2n = 14, BC, Calder & Taylor 35912 (DAO), Taylor & Mulligan (1968). Smelowskia borealis (Greene ) Drury & Rollins 2n = 12, AK, Murie (CAN), Gaiser in Drury & Rollins (1952); Batten 75-455 (ALA), Dawe & Murray (1979); Rice (ALA), Dawe & Murray (1981a); Parker 1734 (ALA), Murray & Kelso (1997). Smelowskia calycina (Stephan ex Willd.) C. A. Mey. var. americana (Regel & Herder) Drury & Rollins 2n = 22, AB, Calder 37261 (DAO), Brockman (unpublished); Mulligan & Mosquin 2675 (DAO) and Mulligan & Crompton 3247 (DAO), both Mulligan (unpublished); Packer 2750 (ALTA), Packer (1968). Smelowskia calycina var. porsildii Drury & Rollins 2n = 22, AK, Packer 1923 (ALTA), Johnson & Packer (1968) as Smelowskia calycina var. integrifolia. The vouchers listed in Dawe & Murray (1979) for Smelowskia calycina subsp. integrifolia var. porsildii, Young 7519 & 7520, with the chromosome number of 2n = 22, are not in ALA as was indicated in the paper (Alan Batten in personal correspondence, May 25, 2001). 2002 Smelowskia media (Drury & Rollins) G. A. Mulligan MULLIGAN: CHROMOSOME NUMBERS OF MUSTARDS 621 2n = 12, AK, Murray 6901 & 6903 (ALA), Dawe & Murray (1981b) as Smelowskia calycina subsp. media. YT, Calder 34367 & 34286 (DAO), Mulligan (unpublished). Smelowskia pyriformis Drury & Rollins 2n = 12, AK, Parker 1256 (ALA), Murray & Kelso (1997). Subularia aquatica var. americana (G. A. Mulligan & Calder) B. Boivin 2n = ca.28, PQ, Lepage 33378 (DAO), Mulligan & Calder (1964). 2n = 30, BC, Calder & Taylor 23653 (DAO), Taylor & Mulligan (1968). Thlaspi arcticum A. Porsild 2n = 14, AK, Murray 6199 (ALA), Dawe & Murray (1981b). Thlaspi arvense L. 2n = 14, ON, Mulligan 1476 (DAO), Mulligan (1957); Pringle 797 (HAM), Pringle (1969); Mulligan & Grainger 4009 (DAO), Mulligan (1984). SK, MacLeod 56 (DAO), Mulligan (1984). AB, Mulligan 1495 (DAO), Mulligan (1957). Acknowledgments I thank the herbarium curators who sent me herbarium chromosome vouchers and provided me with information about specimens under their care. I also want to acknowledge the large contribution to the systematics of the Brassicaceae by Reed C. Rollins that culminated in his Cruciferae of Conti- nental North America (Rollins 1993). I especially want to thank the many authors who deposited vouchers of the plants counted in various herbaria. The identity of the taxa that they counted could not only be checked by me, but also by future research- ers in our continuing attempts to understand the mys- teries of nature. Documents Cited (marked * in text) Mulligan, G. A. 2002. Key to the Brassicaceae (Cruci- ferae) of Canada and Alaska. Electronic publication on the internet site, http://members.rogers.com/mulligan 4520/key/. Literature Cited Bocher, T. W. 1969. Further studies in Arabis holboellii and some allied species. Saertryk Botaniska Tidsskrift 64: 141-161. Dawe, J. C., and D. F. Murray. 1979. IOPB chromosome number reports, LXIII. Taxon 28: 265-268. Dawe, J. C., and D. F. Murray. 1980. IOPB chromosome number reports, LXIX. Taxon 29: 703-730. Dawe, J. C., and D. F. Murray. 198la. IOPB chromo- some number reports, LXX. Taxon 30: 70-72. Dawe, J. C., and D. F. Murray. 1981b. Chromosome numbers of selected Alaskan vascular plants. Canadian Journal of Botany 59: 1373-1381. Drury, J. C., and R. C. Rollins. 1952. The North American representatives of Smelowskia (Cruciferae). Rhodora 54: 85-119. Gervais, C. 1979. Liste annotée de nombres chromosomi- ques de la flore vasculaire du nord-est de I’ Amérique. Le Naturaliste Canadien 106: 451-461. Gervais, C. 1981. Liste annotée de nombres chromo- somiques de la flore vasculaire du nord-est de |’ Amér- ique. II. Le Naturaliste canadien 108: 143-152. Gervais, C., M. Parent, R. Trahan, and S. Plante. 1997. IOPB chromosome data 12. IOPB Newsletter 28: 16. Gervais, C., R. Trahan, and J. Gagnon. 1999. IOPB chromosome data 14. IOPB Newsletter 30: 11. Hedberg, O. 1967. Chromosome numbers of vascular plants from arctic and sub-arctic North America. Arkiv for Botanik 6: 320. Johnson, A. W., and J. G. Packer. 1968. Chromosome numbers in the Flora of Ogotoruk Creek, N.W. Alaska. Botaniska Notiser 121: 403-456. Love, A., and D. Léve. 1975. IOPB chromosome number reports, L. Taxon 24: 673. Love, A., and D. Léve. 1982. IOPB chromosome number reports, LXXIV. Taxon 31: 125-126. Love, A., and J. C. Ritchie. 1966. Chromosome numbers for central northern Canada. Canadian Journal of Botany 44: 429-439. Montgomery, F.H. 1955. Preliminary studies of the genus Dentaria in eastern North America. Rhodora 57: 161-173. Mosquin, T., and D. E. Hayley. 1966. Chromosome numbers and taxonomy of some Canadian arctic plants. Canadian Journal of Botany 44: 1209-1218. Mulligan, G. A. 1957. Chromosome numbers of Can- adian weeds. I. Canadian Journal of Botany 35: 779-789. Mulligan, G. A. 1959. Chromosome numbers of Canadian weeds. II. Canadian Journal of Botany 37: 81—92. Mulligan, G. A. 196la. Chromosome numbers of Can- adian weeds. III. Canadian Journal of Botany 39: 1059-1066. Mulligan, G. A. 1961b. The genus Lepidium in Canada. Madrono 16: 77-90. Mulligan, G. A. 1964. Chromosome numbers of the family Cruciferae. I. Canadian Journal of Botany 42: 1509-1519. Mulligan, G. A. 1965. Chromosome numbers of the family Cruciferae. Il. Canadian Journal of Botany 43: 657-668. Mulligan, G. A. 1966. Chromosome numbers of the family Cruciferae. III. Canadian Journal of Botany 44: 309-319, Mulligan, G. A. 1968. Physaria didymocarpa, P. brassi- coides, and P. floribunda (Cruciferae) and their close relatives. Canadian Journal of Botany 46: 735-740, Mulligan, G. A. 1970a. Cytotaxonomic studies of Draba glabella and its close allies in Canada and Alaska, Canadian Journal of Botany 48: 1431-1437. Mulligan, G. A. 1970b. A new species of Draba in the Kananaskis Range of southwestern Alberta, Canadian Journal of Botany 48; 1897-1898. 622 Mulligan, G. A. 1971a. Cytotaxonomic studies of closely allied Draba cana, D. cinerea and D. groenlandica in Canada and Alaska. Canadian Journal of Botany 49: 89-93. Mulligan, G. A. 1971b. Cytotaxonomic studies of Draba species of Canada and Alaska: D. ventosa, D. ruaxes and D. paysonii. Canadian Journal of Botany 49: 1455-1460. Mulligan, G. A. 1972. Cytotaxonomic studies of Draba species in Canada and Alaska: D. oligosperma and D. incerta. Canadian Journal of Botany 50: 1763-1766. Mulligan, G. A. 1974a. Cytotaxonomic studies of Draba nivalis and its close allies in Canada and Alaska. Can- adian Journal of Botany 52: 1793-1801. Mulligan, G. A. 1974b. Confusion in the names of three Draba species of the arctic: D. adamsii, D. oblongata and D. corymbosa. Canadian Journal of Botany 52: 791-793. Mulligan, G. A. 1975. Draba crassifolia, D. albertina and D. stenoloba in Canada and Alaska. Canadian Journal of Botany 53: 745-751. Mulligan, G. A. 1976. The genus Draba in Canada and Alaska: key and summary. Canadian Journal of Botany 54: 1386-1393. Mulligan, G. A. 1979. Four new species of Draba in northwestern North America. Canadian Journal of Botany 57: 1873-1875. Mulligan, G. A. 1984. Chromosome numbers of some plants native and naturalized in Canada. Le Naturaliste canadien 111: 447-449. Mulligan, G. A. 1995. Synopsis of the genus Arabis (Brassicaceae) in Canada, Alaska and Greenland. Rhodora 97: 108-188. Mulligan, G. A., and J. A. Calder. 1964. The genus Subularia (Cruciferae). Rhodora 66: 127-135. Mulligan, G. A., and W. J. Cody. 1968. Draba norvegica, disjunct to Mackenzie District, NWT. Canadian Journal of Botany 46: 1334-1335. Mulligan, G. A., and W. J. Cody. 1995. New information on the problem of the Asiatic cress, Rorippa crystallina Rollins (Brassicaceae). Canadian Field-Naturalist 109: 111-112. Mulligan, G. A., W. J. Cody, and N. Grainger. 1972. IOPB chromosome number reports, XX XVII. Taxon 21: 498. Mulligan, G. A., and C. Frankton. 1962. Taxonomy of the genus Cardaria with particular reference to the species introduced into North America. Canadian Journal of Botany 40: 1411-1425. Mulligan, G. A., and C. Frankton. 1967. Present status of tall wormseed mustard, Erysimum hieraciifolium, in Canada. Canadian Journal of Botany 45: 755-756. Mulligan, G. A., and D. B. Munro. 1984. Chromosome numbers and sexual compatibility in North American THE CANADIAN FIELD-NATURALIiST Vol. 116 - Rorippa sylvestris (Cruciferae). Canadian Journal of Botany 62: 575-580. Mulligan, G. A., and A. E. Porsild. 1968. A natural first generation hybrid between Rorippa barbareaefolia and R. islandica. Canadian Journal of Botany 46: 1079-1081. Mulligan, G. A., and A. E. Porsild. 1969a. Chromosome numbers of some plants from the unglaciated Yukon plateau, Canada. Canadian Journal of Botany 47: 655-662. Mulligan, G. A., and A. E. Porsild. 1969b. A new species of Lesquerella (Cruciferae) in northwestern Canada. Canadian Journal of Botany 47: 215-216. Mulligan, G. A., and A. E. Porsild. 1970. IOPB chromo- some number reports. XX V. Taxon 19: 111-112. Murray, D. F., and S. Kelso. 1997. Chromosome num- bers and notes on the taxonomy of selected Alaskan vascular plants. Rhodora 99: 33-55. Packer, J. G. 1964. Chromosome numbers and taxonom- ic notes on western Canadian arctic plants. Canadian Journal of Botany 42: 473-494. Packer, J. G. 1968. IOPB chromosome number reports, XVII. Taxon 17: 287. Packer, J. G., and G. D. McPherson. 1974. Chromosome numbers in some vascular plants from Northern Alaska. Canadian Journal of Botany 52: 1095-1099. Packer, J. G., and R. Witkus. 1982. IOPB chromosome number reports, LXXV. Taxon 31: 363. Pringle, J. S. 1969. Documented plant chromosome numbers 1969:1. Sida 3: 350-351. Rollins, R. C. 1941. Monograph study of Arabis in western North America. Rhodora 43: 289-411. Rollins, R. C. 1966. Chromosome numbers of Cruciferae. Contributions to the Gray Herbarium, No. 197: 43-65. Rollins, R. C. 1993. The Cruciferae of Continental North America. Stanford University Press, Stanford, California. 976 pages. Rollins, R. C., and A. E. Shaw. 1973. The genus Lesquerella (Cruciferae) in North America. Harvard University Press, Cambridge, Massachusetts. Taylor, R. L., and R. P. Brockman. 1966. Chromosome numbers of western Canadian plants. Canadian Journal of Botany 44: 1093-1103. Taylor, R. L., and G. A. Mulligan. 1968. Flora of the Queen Charlotte Islands, part 2, cytological aspects of the vascular plants. Queen’s Printer, Ottawa. 148 pages. Taylor, R. L., and S. Taylor. 1977. Chromosome num- bers of vascular plants of British Columbia. Syesis 10: 125-138. Warwick, S., L. D. Black, and J. K. Anderson. 1994. IOPB chromosome data 7. IOPB Newsletter 22: 4—5. Received 26 October 2001 Accepted 12 December 2002 Weedy Introduced Mustards (Brassicaceae) of Canada GERALD A. MULLIGAN Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada Mulligan, Gerald A. 2002. Weedy introduced mustards (Brassicaceae) of Canada, Canadian Field-Naturalist 116(4): 623-631. A list of all 9 of the weedy introduced mustards of Canada is provided, along with information on their introduction, ori- gin, spread, distribution, abundance, and habitats in Canada. Chromosome numbers determined for 41 from Canadian material are included. Key Words. Introduced weedy mustards, Brassicaceae, distributions, habitats, chromosome numbers, Canada Canadian weeds are plants that are prepro- grammed to colonize the many open and disturbed habitats created and maintained by deforestation, agriculture, settlements, and a network of railway lines and roads. Some of these weedy colonizers are native plants that previously only occurred in natu- rally disturbed habitats, but the most aggressive are introduced plants that have evolved as the result of a long association with human activities in the Old World. In addition, many of these introduced colonizers have left their competitors, pests, and par- asites behind. Some of the most successful weedy colonizers in Canada are plants of the mustard fami- ly (Brassicaceae). A few of these, such as Bird Rape (Brassica rapa L.) and wild and cultivated radishes (Raphanus species), have even evolved both weedy and cultivated strains during their long association with humans. The most successful colo- nizing mustards are generally autogamous (produce abundant seed by self-fertilization within the same plant), apomictic (produce abundant seed without the ovaries being fertilized), or have either a strong veg- etative means of reproduction or are common con- taminants in crop seed. Colonization by weeds is completely dependent on the availability of the many disturbed habitats that we have created and continue to maintain. Humans are, in fact, mainly responsible for creating and maintaining the habitats that are necessary for the survival of the very plants that we are most anxious to control or eliminate. This paper provides a listing of the weedy intro- duced mustards of Canada, and information on their origin, introduction, spread, distribution, abundance, and habitats. Included are the chromosome numbers that have been determined from Canadian material. Much of the information included here has resulted from data obtained during two of my recent studies: Key to the Brassicaceae (Cruciferae) of Canada and Alaska (Mulligan 2002*) and Chromosome numbers determined from Canadian and Alaskan material of native and naturalized mustards Brassiceae (Cruciferae) (Mulligan 2002). *see Documents Cited Abbreviations AK: Alaska YT: Yukon NT-M: Northwest Territories- Mackenzie District N-K: Nunavut- Keewatin District N-K: Nunavut- Franklin District NF: Newfoundland and Labrador PE: Prince Edward Island NS: Nova Scotia NB: New Brunswick PQ: Québec ON: Ontario MB: Manitoba SK: Saskatchewan AB: Alberta BC: British Columbia Systematic List Alliaria petiolata (M.Bieb.) Cavara & Grande ( = Alliaria officinalis Andrz. ex M. Bieb.) — Garlic Mustard; Alliaire officinale Self-incompatible biennial. Introduced from Europe for use as a medicinal and salad plant. First collected at Moss Park, Toronto, ON, by J. Fletcher in 1874. Shaded and semi-shaded habitats in NB, PQ, ON, and BC; sporadic except in southern ON. Disagreeable flavor in milk from cows eating garlic mustard. See Cavers et al. (1979). 2n = 42, ON (Mulligan 1984). Alyssum alyssoides (L.) L. — Small Alyssum; Alysson a calices persistants Self-compatible, autogamous, annual or winter annual, introduced from Eurasia. First collected by W. Scott at Lincoln, ON, in 1896. Roadsides, rail- way beds, waste places, ballast, and other disturbed habitats in NF, PQ, ON, MB, AB, and BC; most common in southern ON and southeastern BC, 2n = 32 (Mulligan 1964). Alyssum desertorum Stapt — Yellow Alyssum, Alysson de déserts Self-compatible, autogamous, annual, introduced from eastern Europe and western Asia, First 623 624 collected on a railroad grade at Mortlack, SK, by J. H. Hudson in 1955. Prairie, grain elevators, railway beds, roadsides, and waste places in southern SK, southern AB, and southeastern and southcentral BC. Abundant on sheep rangelands in Rocky Mountain and Great Basin regions of the United States (Rollins 1981). 2n = 32, AB (Mulligan 1964). Alyssum murale Waldst. & Kit. — Yellow-tuft Caespitose perennial. Ornamental, native of Europe, that is a rare escape from cultivation or per- sists at old garden sites in PQ, ON, AB, and BC. First collected at Lambton, ON, by H. N. Racicot in 1947. Alyssum saxitale L. — Golden-tuft Perennial, introduced from Europe. Frequently grown as an ornamental. The only specimen seen was collected in waste ground at Ste-Anne-de-la- Pocatiére, PQ, by L. Cinq-Mars in 1967. It may not persist in nature. Arabidopsis thaliana (L.) Heynh. — Mouse-ear- cress; Arabette des dames Self-compatible, autogamous, annual, introduced from Eurasia. First collected by H. Groh at Agassiz, BC, in 1931. Sporadic in gardens, roadsides, railway beds, and waste places in southern PQ, southern ON, and southern BC. Used extensively in experimental biology. 2n = 10, ON (Mulligan 1984). Arabis caucasica Willd. Wall Rock-cress; — Corbeille d’ argent Self-incompatible, perennial, rock garden plant. Native of Eurasia. First collected by A. Hamel and A. Payette, in tidal debris, at Riviére-du-Loups, PQ, in 1949. Rare garden escape along a roadside in YT, and in cliff faces, tidal rocks, and tidal debris in NB, PQ, and ON. 2n = 16, ON (Mulligan 1964). Arabis glabra (L.) Bernh. (= Turritis glabra L.) — Tower Mustard; Arabette glabre Self-compatible, autogamous, biennial. Both native and introduced from Eurasia. First collected in a sandy field at London, ON, by T. W. Burgess, in 1879. Native on dry slopes in the Cypress Hills of SK and AB, and in the mountainous regions of southwestern AB and southern BC. Probably intro- duced into YT, NT-M, NB, PQ, ON, MB, SK, AB, and BC. Naturalized in sandy roadsides, waste places and field margins throughout its range. 2n = 12, PQ, ON, AB, and BC (Mulligan 1964; Taylor and Mulligan 1968; Mosquin in Mulligan 1995). Arabis hirsuta (L.) Scop. var. hirsuta — Hairy Arabis; Arabette hirsute Biennial or short-lived perennial, native of Europe. The only Canadian specimens were collect- ed by E. Haber and M. J. Schepanek along a road- side in a campground in Glacier National Park, BC, on July 29, 1972, and by V. L. Harms, south of Swift Current Bay of Diefenbaker Lake, SK, on July 21, THE CANADIAN FIELD-NATURALIST Vol. 116 - 1966. Probably introduced from populations known to occur in the United States. Armoracia rusticana P. Gaertn., B. Mey. & Scherb. — Horseradish; Raifort Self-incompatible perennial, introduced from Europe. Thick, deep, branching, woody roots used as source of sauce or relish. Naturalized in PQ as early as 1850 (Rousseau 1968). Roadsides, ditches, waste places, and old garden sites. Sporadic in PE, and from NB to BC. Once established, difficult to eradicate. Barbarea stricta Andrz. — Small Flowered Winter Cress; Barbarée raide Perennial, introduced from Europe. First collected at Gross-Ile, PQ, in St. Lawrence River, in 1944, by R. P. Hanson (Mulligan 1978). Jacques Cayouette has recently seen it at several locations on the island. He also collected it in Comté de Duplessis and Comté de Montmagny, in PQ, in 1979 (Cayouette 1984). It was reported to be at Temagami Forest Reserve, ON (Dorofeev 1998). May be more com- mon than realized because of its confusion with the “typical phase” of Barbarea vulgaris and with Barbarea orthoceras. Barbarea verna (Mill.) Aschers. — Early Winter Cress; Barbariée printaniére Biennial or short-lived perennial, introduced from Europe as a salad plant. First collected at St. Pierre, NF, by L. Arséne in 1900. Roadsides and waste places in NF and on Vancouver Island, BC. Barbarea vulgaris (L.) W. T. Aiton — Yellow Rocket; Barbarée vulgaire Self-incompatible perennial, introduced from Europe, probably as a common impurity in clover, alfalfa, and grass seed. Reproduces both by seed and cauline rosettes. First reported at Montreal in 1821 (Rousseau 1968). NF, PE, NS, NB, PQ, ON, MB, AB, and BC; mostly in non-cultivated fields, roadsides, and waste places. Sometimes used as salad or garnish- ing plant. See MacDonald and Cavers (1991). 2n = 16, ON (Mulligan 1959, 1984). Berteroa incana (L.) DC. — Hoary Alyssum; Bertéroa blanc Annual or winter annual, introduced from Europe. First collected at Wallbridge, ON, by Massey in 1893. Sporadic, but often locally abundant in NS, NB, PQ, ON, MB, SK, and BC. Non-cultivated fields, roadsides, waste places, and settlements; often on poorer soils. 2n = 16, ON (Mulligan 1957). Brassica juncea (L.) Czern. — Indian Mustard; Moutarde d’ Inde Annual, introduced from Europe as a seed con- taminant. Often cultivated for greens. Present in PQ in 1875 (Rousseau 1968) and was collected in a potato field, at Winnipeg, MB, by J. Fletcher in 1896. Cultivated fields, roadsides, and waste places in all provinces from NF to BC. Reaches greatest 2002 abundance in Prairie Provinces. 2n = 36, AB (Mulligan 1959). Brassica napus L. — Rape; Navet Annual, introduced from Eurasia. Sporadic escape from cultivation. First collected at St. John, NB, by G. F. Matthew in 1874. Rare plant in cultivated and abandoned fields, roadsides, railways, and waste places in NT-M and every province from NF to BC. Brassica nigra (L.) W. D. J. Koch — Black Mustard; Moutarde noire Annual, introduced from Europe. An escape from cultivation and a seed contaminant. Naturalized in Canada since 1863 (Groh and Frankton 1946). Sporadic in cultivated fields, gardens, waste places, and roadsides in NF, NS, NB, PQ, ON, AB, and BC. Brassica oleraceae L. — A great number of agricul- tural and horticultural varieties Annual, introduced from Eurasia. Many cultivated varieties escape from cultivation but do not persist in Canada. Brassica rapa L. (= Brassica campestris L.) — Bird Rape; Moutarde des oiseaux Annual or winter annual, introduced from Eurasia. Has oil seed, vegetable, salad plant, and weed phases. Mostly introduced into fields as a contaminant in crop seed. In NS as early as 1829 (Rousseau 1968).Cultivated fields, roadsides, and waste places in YT, NT-M, and in all provinces from NF to BC. Most common in the Maritime Provinces, maritime areas of PQ, and the Fraser Valley and Vancouver Island of BC. 2n = 20, ON and BC (Mulligan 1959; Taylor and Mulligan 1968). Bunias orientalis L. — Turkish Rocket; Bunias d’ Orient Perennial, introduced from Eurasia. Very spo- radic in waste land and along roadsides in NS, NB, southern PQ, and southern BC. Does not persist except at a few locations on Vancouver Island and in the Fraser Valley of BC. Collected at Botanie, BC, by W. B. Anderson in 1927 and at the same site by F. Lomer in 1995. Cakile edentula (Bigelow) Hook. var. edentula — Sea Rocket; Caquillier édentulé Annual coastal strand herb, native to east coast but introduced on the west coast. In NF, PE, NS, NB, PQ, and BC. 2n = 18, PQ and BC (Mulligan 1964; Taylor and Mulligan 1968). Cakile maritima Scop. — Sea Rocket; Caquillier maritime Annual seashore strand herb, introduced from Europe. West coast, from Queen Charlotte Islands, BC, to Baja California. Sporadic along east and Gulf coasts of the United States (Rollins 1981). Camelina alyssum (Mill.) Thell. (= Camelina parodii Ibarra & La Porte) — Flat-seeded False Flax; Caméline alysson MULLIGAN: WEEDY INTRODUCED MUSTARDS OF CANADA 625 Annual, introduced from Europe as an impurity in crop seed. First collected at Middle Lake, SK, by W. W. Robins in 1910. Infrequent in cultivated fields, roadsides and waste places in the southern parts of MB, SK, and AB. Camelina microcarpa Andrz. ex DC. — Small- seeded False Flax; Caméline 4 petits fruits Self-compatible, autogamous, annual, introduced from Eurasia. Contaminant in flax and grain seed. First collected at Snelgrove, ON, by J. White in 1897. In all provinces, but most common in the Prairie Provinces. Open prairie, cultivated fields, roadsides, railway beds, and waste places. 2n = 40, AB and BC (Mulligan 1957, 1984). Camelina sativa (L.) Crantz — Large-seeded False Flax; Caméline cultivée Self-compatible, autogamous, annual, introduced from Eurasia. Contaminant in flax and grain seed. First collected at Fish Creek, AB, by J. Fowler in 1894. In YT (rare), NT-M, and from PQ to BC. Less common than small-seeded false flax. Prairies, culti- vated fields, around grain elevators, roadsides, rail- ways, and waste places. 2n = 40, AB and BC (R. J. Moore, unpublished). Capsella bursa-pastoris (L.) Medik. — Shepherd’s Purse; Bourse-a-pasteur Autogamous, self-compatible, annual or winter annual, introduced from Europe. Recorded in New England as early as 1672 (Rousseau 1968) and in Nova Scotia as early as 1829 (Groh 1945). In YT, NT-M, N-F, and all provinces from NF to BC. Common plant in cultivated fields, pastures, mead- ows, roadsides, railway beds, and waste places, in all settled areas of Canada. 2n = 32, ON, SK and BC (Mulligan 1957, 1984; Taylor and Mulligan 1968). Cardamine flexuosa With. — Wavy Bitter Cress; Cardamine flexueuse Biennial or short-lived perennial, introduced from Europe. First collected in a park at St. John’s, NF, by A. M. Ayre in 1927. Rare in wet springy banks and thickets in NF. Cardamine hirsuta L. — Hoary Bitter Cress; Cardamine hirsute Autogamous, self-compatible, annual, introduced from Europe. First collected at Vancouver, BC, by J. R. Anderson in 1896. Rare in ON, and common on Vancouver Island and in the Fraser Valley of BC. Old fields, gardens, ditches, roadsides, and waste places, usually in moist soil. 2n = 16, BC (Mulligan 1965). Cardamine impatiens L. — Narrow Leaved Bitter Cress; Cardamine impatiente Annual or biennial herb introduced from Europe. Only specimen seen was collected from a deciduous woods on St. Lawrence Starch Works property, Port Credit, ON, by J. M. Weber in 1980. It is sporadic and uncommon in the United States (Rollins 1981). 626 Cardamine pratensis L. — Cuckoo Flower; Cardamine des prés Perennial. In YT, NT-M, N-K, N-F, NS, and from NB to BC. Plants in southern part of its range are mostly rare ornamental escapes of Eurasiatic ori- gin. Plants in the northern part of its range, in Canada, are probably native. Cardaria chalepensis (L.) Hand.-Mazz. — Lens- podded Hoary Cress; Cranson rampant Strongly rhizomatous perennial, introduced into Canada and the United States in alfalfa seed import- ed from Turkestan about 1911 or 1912. First collect- ed in the United States at Fargo, North Dakota, in 1912, and in Canada at Grande Prairie, AB, in 1926. In cultivated and non-cultivated habitats from ON to BC. Most abundant in irrigated areas of western Canada and the western United States. The three hoary cresses, found in Canada, now survive and spread primarily by extremely persistent vertical and horizontal roots. This is the most aggressive of the three hoary cresses. Fields, ditches, roadsides, pas- tures, and waste places; often on saline soils. See Mulligan and Findlay (1974). 2n = 80, SK, AB and BC (Mulligan and Frankton 1962). Cardaria draba (L.) Desv. — Heart-podded Hoary Cress; Cranson dravier Rhizomatous perennial, introduced from Europe and Western Asia. First collected in the United States at Long Island, New York, in 1862, and in Canada at Barrie, ON, in 1878. Present in NS, and from ON to BC. It is most troublesome in southern MB and southern AB. Cultivated fields, hayfields, pastures, irrigation ditches, roadsides, and waste places. Least weedy of the hoary cresses. See Mulligan and Findlay (1974). 2n = 64, AB (Mulligan and Frankton 1962). Cardaria pubescens (C. A. Mey.) Jarmol. Globe- podded Hoary Cress; Cranson velu Rhizomatous perennial introduced into Canada and the United States in alfalfa seed imported from Turkestan in 1911 or 1912. In MB, SK, AB, and BC. Often found in the same fields as lens-podded hoary cress, but not nearly as aggressive. First collected in the United States at Fargo, North Dakota, in 1912, and in Canada at Grande Prairie, AB, in 1926. In cultivated and non-cultivated fields, especially in irrigated areas where saline soils are present. See Mulligan and Findlay (1974). 2n = 16, SK and AB (Mulligan and Frankton 1962). Chorispora tenella (Pallas) DC. — Blue Mustard; Chorispora fluet Annual or winter annual, introduced from Asia. First collected at Penticton, BC, by J. W. Eastham in 1940. Very rare and often not persisting. In the southern parts of SK, AB, and BC. Very abundant in rangelands of the United States (Rollins 1981). THE CANADIAN FIELD-NATURALIST Vol. 116 - Conringia orientalis (L.) Dumort. — Hare’s-ear Mustard; Vélar d’ Orient Autogamous annual or winter annual, introduced from Eurasia. First collected at Bass River, NB, by J. Fowler in 1872. In all provinces from NF to BC. Reaches greatest abundance in the Prairie Provinces, especially SK. Grain fields, disturbed prairie, pas- tures, roadsides, railways, and waste places. Coronopus didymus (L.) Sm. — Lesser Swine Cress; Corne-de-cerf didyme Annual or winter annual, introduced from Asia or South America. First collected on a wharf at Gaspé Basin, PQ, by J. Bell in 1862 and at North Sidney, NS, by T. W. Burgess in 1883. Rare in ballast, road- sides, waste places, and in old fields along the coasts of NF, NS, NB, PQ, and southern BC. Coronopus squamatus (Forssk.) Asch. — Creeping Wart Cress; Corne-de-cerf écailleuse Biennial or short-lived perennial, introduced from Europe. First collected on a wharf at Gaspé Basin, PQ, by J. Bell in 1862 and at St. John, NB, by J. Fowler in 1877. Rare in coastal areas of NS, NB, and PQ, and on lake shores in ON. Descurainia sophia (L.) Webb. — Flixweed; Sagesse-des-chirurgiens Autogamous, self-compatible, annual or winter annual, introduced from Eurasia. At Montreal, PQ, in 1821; probably reaching Canada with early French settlers (Rousseau 1968). First collected, in the prairies, at Plum Coulee, MB, by J. Fletcher in 1907, and at Antler, SK, by E. L. Stonehouse in the same year. In YT, NT-M, and from PE to BC. Most common in the Prairie Provinces, especially in cereal crops. Sporadic elsewhere. In disturbed grasslands, hay fields, grain fields, roadsides, and waste places. See Best (1977). 2n = 28, SK, AB and BC (Mulligan 1961b, 1984; Taylor and Mulligan 1968). Diplotaxis erucoides (L.) DC. — White Rocket; Diplotaxe fausse-roquett Annual or winter annual, native of Europe. In Canada, only known from PQ; Gaspé-Nord in 1904 and Laval in 1961 (Rousseau 1968). On ballast in New York City and in Camden, New Jersey, prior to 1880 (Rollins 1981). Very rare plant on ballast and in waste places. May not persist. Diplotaxis muralis (L.) DC. — Sandrocket; Diplotaxe des murs Autogamous, annual or winter annual, introduced from Europe. First collected on ship ballast at North Sydney, NS, and at Pictou, NS, by J. Macoun in 1883. Sporadic, but locally abundant, from PE to AB. In disturbed grassland, gardens, railway beds, and waste places, and around buildings. Diplotaxis tenuifolia (L.) DC. — Wall-rocket; diplotaxe a feuilles ténues Perennial, introduced from Europe. First collected at St. John, NB, by J. Fowler in 1877. NS, NB, PQ, 2002 ON, and BC. Most common in southern ON; spo- radic elsewhere. In waste places, ballast, railway beds, and along roadsides. 2n = 22, ON (Mulligan 1959). Draba nemorosa L. Wood — Whitlow-grass; Drave des bois Native, autogamous, annual or winter annual. In YT, NT-M, PQ, and from PQ to BC. Introduced in Eastern Canada and native elsewhere. Has colonized a wide variety of artificially disturbed habitats both within and outside of its native range. 2n = 16, ON, MB and AB (Mulligan 1966, 1975; Packer 1964; Love and Love 1982). Draba verna L. — Whitlow-grass; Drave printaniére Autogamous, annual or winter annual, introduced from Europe. First collected at Quebec, PQ, by J. Bell in 1865. NB, and southern parts of PQ, ON, AB, and BC. In disturbed grasslands, railway beds, roadsides, and waste places, especially on light soils. 2n = 39, BC (Mulligan 1966). Eruca vesicaria (L.) Cav. subsp. sativa (Mill.) Thell. — Garden-rocket; Roquette des jardins Annual, introduced from Eurasia. Grown for salad purposes. First collected at Grenfell, SK, by Deacon in 1908. Early introductions were from contaminants in alfalfa seed (Groh 1944). Sporadic in waste places, cultivated fields, and roadsides in PQ, and southern parts of ON, SK, AB, and BC. Erucastrum gallicum (Willd.) O. E. Schulz — Dog Mustard; Moutarde des chiens Autogamous annual or winter annual, introduced from Eurasia. First collected at Emerson, MB, by H. Groh in 1943. Early collections were along railway beds, near grain elevators, and at grain shipping ter- minals. In cultivated fields, roadsides, railway beds, and waste places in NT-M, and from NF to BC. See Warwick and Wall (1998). 2n = 30, ON (Mulligan 1957, 1984). Erysimum cheiri (L.) Crantz Wallflower Perennial, introduced from Europe, grown as an ornamental. An infrequent garden escape near Whitehorse, YT, at Ste-Foy, PQ, and on southeast Vancouver Island, B.C. — Common Erysimum cheiranthoides L. — Wormseed Mustard; Vélar fausse-giroflée Autogamous annual or winter annual, introduced from Eurasia. First recorded in the United States, in Virginia, by Pursh in 1814 and, in Canada, by Hooker in 1830 (Rousseau 1968). Present in YT, NT-M, and in all provinces from NF to BC. Ina wide range of disturbed habitats, but it is usually sparse at any single location. 2n = 16, ON and BC (Mulligan 1957, 1984; Taylor and Mulligan 1968). Erysimum hieraciifolium L. Tall — Wormseed Mustard; Vélar a feuilles d’éperviere MULLIGAN: WEEDY INTRODUCED MUSTARDS OF CANADA 627 Apomictic biennial or short-lived perennial, intro- duced from Europe. First seen at Ottawa, ON, in 1941 and collected at Apple Hill, ON, by E. G. Anderson in 1946. In NS, NB, PQ, ON, and SK. Reaches greatest abundance, and is spreading rapid- ly, in roadsides, gravel pits, pastures, and hay fields in eastern ON. 2n = 48, meiosis irregular, ON (Mulligan and Frankton 1967). Erysimum repandum L. — Treacle Mustard; Vélar étale Annual or winter annual, introduced from Eurasia. First collected on Pelee Island, ON, by W. Botham in 1938. A rare mustard, present only at a few sites in waste places, railway beds, and roadsides in Essex and Kent Counties in southern ON, and in the Fraser Valley of BC. Hesperis matronalis L. — Dame’s Rocket; Julienne des dames Perennial, introduced from Europe as an ornamen- tal plant. Naturalized in PQ as early as 1862 (Rousseau 1968). Sporadic in old garden sites, road- sides, and waste places in all provinces. 2n = 24, ON (Mulligan 1984). Hutchinsia procumbens (L.) Desv. (= Horungia procumbens (L.) Hayek) — Oval Shepherd’s Purse; Hutchinsie couchée Annual herb, introduced from Eurasia. First collect- ed at Cache Creek, BC, by J. Macoun in 1875. Occurs at a few sites, on moist alkaline flats and seashores, in NF, MB, southern SK, and southern BC. Iberis amara L. — Rocket Candytuff; Iberis amer Annual, introduced from Europe as an ornamental. First collected at Lindsay, ON, in 1903 by W. Scott. A rare escape, in roadsides and waste places, in NS, and ON. Iberis umbellata L. — Globe Candytuff; Iberis a ombelles Annual, introduced from Europe as an ornamental. A rare escape from cultivation that is sometimes found in roadsides and waste places in PE, NS, and ra. Isatis tinctoria L. Dyers — Woad; Pastel des teinturiers Biennial, formerly cultivated for its blue dye. Sparingly escaped from cultivation in NF, PQ, ON, and BC. First collected at Toronto, ON, by H. Groh in 1924. Only persists at a few locations in roadsides and waste places, in the Fraser Valley and on Vancouver Island of BC. Weedy Native Pepper-grasses; Lepidium species Lépidies Four, autogamous, annual or biennial pepper- grasses are native in some parts of Canada, but have colonized a wide range of artificially disturbed habitats both within and outside of their native ranges. 628 Lepidium bourgeauanum Thell. — Bourgeau’s Pepper-grass. A biennial. In YT, NT-M, NF, NB, and PQ to BC. Native to Prairie Provinces but introduced else- where. 2n = 32, ON and BC (Mulligan 1961a). Lepidium densiflorum Schrad. — Common Pepper- grass; Lépidie densiflore. An annual. In YT, NT-M, and from NF to BC. Native to Prairie Provinces and interior of BC and introduced elsewhere. 2n = 32, ON, SK and BC (Mulligan 1961a,b and 1984). Lepidium ramosissimum A.Nelson — Western Pepper-grass. A biennial. In NT-M, ON, MB, SK, AB, and BC. Native of Prairie Provinces and introduced else- where. 2n = 64, NT-M, SK and AB (Mulligan 1961a). Lepidium virginicum L. — Poor-man’s Pepper-grass; Lépidie de Virginie. An annual. In NF, PE, NS, PQ, ON and south- western BC. Native to Fraser Valley and adjacent Vancouver Island in BC, but introduced in the east from further south in the United States. 2n = 32, ON and BC (Mulligan 1961a). Lepidium aucheri Boiss. Introduced from Europe. Only specimen seen was collected on Bull grounds, Toronto, ON, by W. Scott on August 20, 1904. Lepidium campestre (L.) R.Br. — Field Pepper- grass; Lépidie des champs Autogamous annual or winter annual, introduced from Europe. First collected at Hamilton, ON, by J. M. Buchan in 1870 (Rousseau 1968). In all provinces of Canada; cultivated fields, roadsides, waste places, and railroad beds. Most common and abundant in the southern parts of PQ, ON, and BC; sporadic elsewhere. 2n = 16, ON, SK and BC (Mulligan 1957; Taylor and Mulligan 1968). Lepidium heterophylum (DC.) Benth. — Smith’s Pepper-grass; Lépidie a feuilles dissemblables Perennial, introduced from Europe. Well estab- lished along roadsides and in waste places on Vancouver Island, B.C. First collected at Courtenay, BC, by W. E. Molyneux in 1955. Lepidium latifolium L. — Perennial Pepper-grass; Lépidie a feuilles larges Perennial, with subterranean rhizomes, introduced from Europe. Only known in Canada at Quebec, PQ, near Lethbridge, AB, and at Wycliffe, in the East Kootenay region of BC. It has persisted and spread locally at all three locations. It was first collected, in the railroad shop yard at Quebec, PQ, by Br. Marie- Anselm in 1931 and was discovered, in irrigated land, near Lethbridge, AB, by K. W. Hill in 1940. 2n = 24, AB (Mulligan 1957). THE CANADIAN FIELD-NATURALIST Ses tt(iti‘a” Vol. 116 - Lepidium oxycarpum Torr. & A. Gray. — Forked Pepper-grass Annual, introduced from the United States. Only Canadian specimen seen was collected at Cadboro Bay, Vancouver Island, BC, by J. Macoun in 1893. Lepidium perfoliatum L. — Clasping-Leaved Pepper-grass; Lépidie perfoliée Autogamous annual, introduced from Europe. Often breaking off at base to act as a tumbleweed for seed dispersal. First collected at Osoyoos and Cranbrook, BC, by H. Groh in 1931. In southwest PQ, and in the southern parts of ON, MB, SK, AB, and BC. Most common in cultivated fields, over- grazed rangeland, roadsides, and waste places in the drier regions of the Prairie Provinces and inland valleys of BC. Lepidium ruderale L. — Roadside Pepper-grass; Passerage des decombres Autogamous annual or winter annual, introduced from Eurasia. First collected by J. Fowler at Bass River, NB, in 1868. Sporadic in roadsides, waste places and settlements from NF to SK. Lepidium sativum L. — Garden Cress; Lépidie cultivée Annual, introduced from Europe as a salad green. First collected at Ottawa, ON, by W. Scott in 1890. Sparingly escaped to roadsides, waste places and set- tlements in YT, NT-M, and from PE to BC. Lobularia maritima (L.) Desv. — Sweet Alyssum; Allysson maritime Perennial ornamental, native to the Mediterranean region, that normally acts as an annual under our conditions. First collected at Tring Junction, PQ, by H. Groh in 1927. Has been collected in NS, southern PQ and ON, and on Vancouver Island BC. It does not appear to persist except on Vancouver Island. Lunaria annua L. — Honesty; Monnaie-du-pape Annual or biennial, native of Europe, grown for ornamental purposes. First collected at Revelstoke, BC, by H. Groh in 1939. Has been collected in old garden sites, roadsides, and waste places in NS, PQ, ON, MB, and BC. Does not persist. Malcolmia maritima (L.) W. T. Aiton — Virginia Stock; Malcolmie maritime Annual ornamental, introduced from Europe. First collected as a garden escape, at Durham, ON, by W. Scott, in 1897. It has not been collected since that time. Matthiola longipetala (Vent.) DC. (= Matthiola bicornis (Sm.) DC.) — Evening Stock Annual or winter annual ornamental, native of Eurasia. First collected at Saskatoon, SK, by H. Groh in 1935. Collected, at a few sites, in settkements and roadsides, in ON, SK, and AB. Does not persist. 2002 Myagrum perfoliatum L — Mitre Cress; Myagrum perfolie Annual, native of Eurasia. Sometimes grown as an ornamental. Collected at Gallow’s Hill, PQ, by Brodie in 1895. The only other record is based on a specimen collected in the 1940s at Rigaud, PQ (Groh and Frankton 1946). There is no evidence that it can persist in nature. Nasturtium crystallinum (Rollins) G. A. Mulligan — Asiatic Cress Perennial with a very thick root. Probably intro- duced from Asia and utilized to produce a condi- ment. First collected by Thieret and Reich at mile 39, Yellowknife Highway, 26 miles northeast of Fort Providence, NT-M, in 1961. This is the only known location for this mustard in North America. Another specimen was collected, from the large patch grow- ing at this site, by W. J. Cody in 1965. This is a loca- tion that was formerly farmed by a religious commu- nity. 2n = 32, NT-M (Mulligan and Cody 1995). Nasturtium microphyllum Boenn. ex Rchb. — Northern Water-cress; Cresson de fontaine Perennial, aquatic to semi-aquatic, rooting at nodes. Introduced from Europe as a salad plant and has spread from deliberate plantings into cold streams. First collected in Canada at London, ON, by T. W. Burgess in 1879. Collected intermittently in cool water habitats in NF, PE, NB, PQ, ON, MB, and BC. Most common in southern ON. 2n = 64, PE and ON (Mulligan 1984). Nasturtium officinale W.T. Aiton — Southern Water-cress; Cresson de fontaine Perennial, aquatic to semi-aquatic, rooting at nodes. Introduced from Europe as a salad plant and has spread from deliberate plantings in cold streams. First collected in Canada at London, ON, by T. W. Burgess in 1878. Intermittently established in cool water in NF, NS, NB, PQ, ON, MB, AB, and BC. Most common in southwestern BC. Frequently confused with northern water-cress. Nasturtium offic- inale, a diploid (2 n = 32), is the most common water-cress in the United States, whereas Nasturtium microphyllum, a tetraploid (2 n = 64), is most common in Canada (Green 1962). Neslia paniculata (L.) Desv. — Ball Mustard; Neslie paniculée Self-compatible, autogamous, annual or winter annual, introduced from Europe. Pods, that do not open when ripe, are frequently an impurity in grain seed. First collected on a railroad embankment at Silver City, AB, by J. Macoun in 1885. In YT, NT- M, and all provinces from NF to BC. Common in grain and other cultivated fields in Western Canada, particularly in the Peace River Districts of AB and BC. Sporadic elsewhere in roadsides, railway embankments, waste places, and near grain eleva- tors. 2n = 14, ON and BC (Mulligan 1957; Taylor and Mulligan 1968). MULLIGAN: WEEDY INTRODUCED MUSTARDS OF CANADA 629 Raphanus raphanistrum L. — Wild Radish; Radis sauvage Annual or winter annual, introduced from Europe. Present in Nova Scotia as early as 1829 (Rousseau 1968). In all provinces from NF to BC. Very abun- dant in cultivated fields along the Atlantic seaboard and in the Fraser Valley and Vancouver Island of BC. Sporadically distributed elsewhere, mostly due to contaminants in grain shipments. 2n = 18, NB (Mulligan 1961b). Raphanus sativus L. — Radish; Radis Annual, native of Mediterranean region, that is a sporadic escape from cultivation. Collected in NS, NB, PQ, ON, MB, SK, and BC, but does not persist, except in the Fraser Valley and Vancouver Island of BC. Sometimes hybridizes with wild radish in the Fraser Valley. Rapistrum perenne (L.) All. — Steppe Cabbage; Rapistre vivace Deep rooted perennial, introduced from Europe. First collected, at its only known location, at Grenfell, SK, in 1928 (Groh 1944). It has persisted and spread locally in roadsides, waste places, and cultivated fields. Rapistrum rugosum (L.) All. — Turnip Weed; Rapistre rugueux Annual, introduced from Europe. First collected at Montreal, PQ, by J. M. Wallace in 1893. Collected at a few widely separated sites, in way- sides, and waste places in PQ and ON. Does not seem to persist. Rorippa amphibia (L.) Besser — Amphibious Water Cress; Cresson amphibie Aquatic to semi-aquatic perennial, introduced from Europe. Rhizomes are believed to have been introduced at the port of Montreal, PQ, in ballast from early European ships. It occurs along the St. Lawrence River from Montreal to Quebec, PQ, and along the Richelieu River at St-Ours, PQ (see Rousseau 1968). Rorippa austriaca (Crantz) Besser — Austrian Yellow Cress; Cresson d’ Autriche Rhizomatous perennial, introduced from Europe. First collected at Greenstreet, SK, by Patmore in 1932. In a few widely scattered localities in MB, SK, and AB. Is in pastures, cropland, and waste places, usually where moisture is plentiful. Rorippa calycina (Engelm.) Rydb. Rhizomatous perennial, probably introduced into Canada, from its native range further south in the United States, by migrating waterfowl. Only known in Canada from the mouth of the Anderson River, NT-M (Mulligan and Porsild 1966). First collected by T. W. Barry in 1962, and in the same area by G. W. Scotter in 1965. 630 Rorippa sylvestris (L.) Besser — Creeping Yellow Cress; Rorippa sylvestre Rhizomatous perennial, introduced from Europe. Self-incompatible, rarely setting seed pods in nature. Spread mainly by root fragments. First collected at Ball’s Mills, Lincoln County, ON, by McCalla in 1897. In all provinces from NF to BC. Present in gardens, ditches, disturbed stream banks, moist meadows, plant nurseries, and waste places. 2n = 32, PQ and ON; 2n = 40, ON and BC; 2n = 48, NB, PQ, ON, AB and BC (Mulligan and Munro 1984). Sinapis alba L. — White Mustard; Moutarde blanche Annual, native of Mediterranean region. Cultivated for the manufacture of mustard from seeds, and to a lesser extent for greens. Growing wild in Canada as early as 1860 (Groh and Frankton 1946). Sporadic in fields, disturbed prairie, road- sides, railway beds, and waste places, in PE and from PQ to BC. Often does not persist. Sinapis arvensis L. — Wild Mustard; Moutarde des champs Self-incompatible annual, introduced from the Mediterranean region. Common in fields of Albany, New York, as early as 1748, in NS in 1829 and by 1840 had reached the western and northern part of the state of New York and Lower Canada. In Fort Garry, MB, in 1860 and is now through all settled areas of Canada (Mulligan and Bailey 1975). Now in YT, NT-M, and all provinces. Jacobson et al. (1988) presented some evidence that charred seeds uncov- ered at excavations of abandoned native sites in the northeastern United States and dated at 8000 B.P. (before present), were wild mustard seeds. Rollins (1993) found their evidence unconvincing, and I agree with him. Very common in grain fields, other cultivated fields, and waysides in the Canadian prairie region, but less common elsewhere. It is a common contaminant in grain seed. Poisoning and death have occurred when large quantities of screen- ings were fed to animals (Mulligan and Munro 1990). See Warwick et al. (2000). 2n = 18, ON, MB, SK, AB and BC (Taylor and Mulligan 1968; Mulligan 1984; Warwick et al. 1994). Sisymbrium altissimum L. — Tumble Mustard; Sisymbre élevé Autogamous annual or winter annual, introduced from Eurasia. First collected at Silver City, AB, by J. Macoun in 1885. Often breaking off at the base, act- ing as a tumbleweed for seed dispersal. In YT, NT- M, and in all provinces from NF to BC. Most abun- dant in grain fields and grasslands of the prairies of Western Canada. Mostly in roadsides, railway beds, and waste places elsewhere in Canada. 2n = 14, ON, SK and BC (Mulligan 1961b, 1984; Taylor and Mulligan 1968). THE CANADIAN FIELD-NATURALIST | aT aC Vol. 116 - Sisymbrium loesellii L. — Tall Hedge Mustard; Sisymbre de Loesel Autogamous annual or winter annual, introduced from Europe. First collected by N. Criddle in SK in 1929. Occurs from NB to BC. Most common in grain fields and other cultivated fields in the Prairie Provinces; sporadic in roadsides and waste places elsewhere. 2n = 14, AB (Mulligan 1957). Sisymbrium officinale (L.) Scop. — Hedge Mustard; Sisymbre officinal Annual, introduced from Europe. Seen at Prescott, ON, by B. Billings in 1862 (Rousseau 1968). Occurs in all provinces, except SK. Primarily a garden and waste place weed. It is often found near farm build- ings. 2n = 14, BC (Taylor and Mulligan 1968). Sisymbrium orientale L. — Eastern Tumble Mustard Annual herb, introduced from Eurasia. Rare in mesic to dry fields, roadsides, and waste places at Vancouver, BC (Douglas et al. 1998). Teesdalia nudicaulis (L.) R.Br. — Shepherd’s Cress; Teesdalie 4 tige nue Annual or winter annual, introduced from Europe. Sporadically naturalized in disturbed habitats on southern Vancouver Island and in the Fraser Valley of BC. First collected in Locarno Park, Vancouver, BC, by J. Eastham in 1938, and it continues to per- sist at that site. Thlaspi arvense L. — Stinkweed; Tabouret des champs Autogamous annual or winter annual, introduced from Europe. It was a common weed around Detroit as early as 1818 and, in Canada, at Fort Garry, MB, in 1865 (Best and McIntyre 1975). In YT, NT-M, and all provinces from NF to BC. In grain fields, hay fields, gardens, roadsides, and railway beds. When grazed, produces an off-flavor in dairy products (Frankton and Mulligan 1987). 2n = 14, ON, SK, and AB (Mulligan 1957, 1984; Pringle 1969). Acknowledgments The publications, Weeds of the Cruciferae (Brassicaceae) in North America (Rollins 1981) and Histoire, habitat et distribution de 220 plantes intro- duites au Québec (Rousseau 1968) have been espe- cially useful to me. The common names are mostly from Common and scientific names of weeds in Canada/Noms populaires et scientifiques des plantes nuisible du Canada (Darbyshire et al. 2000). I am also indebted to the many people who documented the first sightings and later spread of our introduced mustards in their publications or by depositing specimens in various herbaria. Documents Cited (marked * in the text) Mulligan, G. A. 2002. Key to the Brassicaceae (Cruciferae) of Canada and Alaska. Electronic publi- cation on the web site, Attp://members.rogers.com/ mulligan4520/keyl. 2002 Literature Cited Best, K. F. 1977. The biology of Canadian weeds. 22. Descurainia sophia (L.) Webb. Canadian Journal of Plant Science 57: 499-507. Best, K. F., and G. I. McIntyre. 1975. The biology of Canadian weeds. 9. Thlaspi arvense L. Canadian Journal of Plant Science 55: 279-292. Cavers, P. B., M. I. Heagy, and R. F. Kokron. 1979. The biology of Canadian weeds. 35. Alliaria petiolata (M. Bieb.) Cavara & Grande. Canadian Journal of Plant Science 59: 217-229. Cayouette, J. 1984. Nouvelles stations du Barbarea stricta Andrz. au Québec. Le Naturaliste Canadien 111: 207-209. Darbyshire, S. J., M. Favreau, and M. Murray. 2000. Common and scientific names of weeds in Canada/ Noms populaires et scientifiques des plantes du Canada. Research Branch, Agriculture and Agri-Food Canada Publication 1397/B. 132 pages. Dorofeev, V. I. 1998. The four new species of Brassicaceae in North America. Botanichiskii Zhurnal 83(9): 133-135. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998. Illustrated Flora of British Columbia, Volume 2. Ministry of Environment, Lands and Parks, Victoria, British Columbia. 401 pages. Frankton, C., and G. A. Mulligan. 1987. Weeds of Canada. NC Press Ltd., Toronto, Ontario. 217 pages. Green, P. S. 1962. Watercress in the New World. Rhodora 64: 32-43. Groh, H. 1944. Canadian weed survey, third annual report. Canada Department of Agriculture, Ottawa, Ontario. 70 pages. Groh, H. 1945. Canadian weed survey, fourth annual report. Canada Department of Agriculture, Ottawa, Ontario. 56 pages. Groh, H., and C. Frankton. 1946. Canadian weed survey, fifth annual report. Canada Department of Agriculture, Ottawa, Ontario. 86 pages. Jacobson, H. A., J. B. Petersen, and D. E. Putnam. 1988. Evidence of pre-Columbian Brassica in northeastern U.S. Rhodora 90: 355-362. Love, A., and D. Léve. 1982. IOPB chromosome number reports, LXXIV. Taxon 31: 125-126. MacDonald, M. A., and P. B. Cavers. 1991. The biology of Canadian weeds. 97. Barbarea vulgaris R.Br. Canadian Journal of Plant Science 71: 49-166. Mulligan, G. A. 1957. Chromosome numbers of Canadian weeds. I. Canadian Journal of Botany 35: 779-789. Mulligan, G. A. 1959. Chromosome numbers of Canadian weeds. II. Canadian Journal of Botany 37: 81-92. Mulligan, G. A. 1961a. The genus Lepidium in Canada. Madrono 16: 77-90. Mulligan, G. A. 1961b. Chromosome numbers of Canadian weeds. III. Canadian Journal of Botany 39: 1059-1066. Mulligan, G. A. 1964. Chromosome numbers of the family Cruciferae. I. Canadian Journal of Botany 42: 1509-1519. Mulligan, G. A. 1965. Chromosome numbers of the family Cruciferae. II. Canadian Journal of Botany 43: 657-668. Mulligan, G. A. 1966. Chromosome numbers of the family Cruciferae. II. Canadian Journal of Botany 44: 309-319. Mulligan, G. A. 1975. Draba crassifolia, D. albertina, D. nemorosa, and D. stenoloba in Canada and Alaska. Canadian Journal of Botany 53: 745-751. MULLIGAN: WEEDY INTRODUCED MUSTARDS OF CANADA 631 Mulligan, G. A. 1978. Barbarea stricta Andrz., a new intro- duction to Quebec. Le Naturaliste canadien 105: 298-299. Mulligan, G. A. 1984. Chromosome numbers of some plants native and naturalized in Canada. Le Naturaliste canadien 111: 447-449. Mulligan, G. A. 1995. Synopsis of the genus Arabis (Brassicaceae) in Canada, Alaska, and Greenland. Rhodora 97: 108-188. Mulligan, G. A. 2002. Chromosome numbers determined from Canadian and Alaskan materal of native and natu- ralized mustards, Brassicaceae (Cruciferae). Canadian Field-Naturalist 116(4): 611-622. Mulligan, G. A., and L. G. Bailey. 1975. The biology of Canadian weeds. 8. Sinapis arvensis L. Canadian Journal of Plant Science 58: 171-183. Mulligan, G. A., and W. J. Cody. 1995. New information on the problem of Asiatic cress, Rorippa crystallina Rollins (Brassicaceae). Canadian Field-Naturalist 109: 111-112. Mulligan, G. A., and J. N. Findlay. 1974. The biology of Canadian weeds. 3. Cardaria draba, C. chalepensis, and C. pubescens. Canadian Journal of Plant Science 54: 149-160. Mulligan, G. A., and C. Frankton. 1962. Taxonomy of the genus Cardaria with particular reference to the species introduced into North America. Canadian Journal of Botany 40: 1411-1425. Mulligan, G. A., and C. Frankton. 1967. Present status of tall wormseed mustard, Erysimum hieraciifolium, in Canada. Canadian Journal of Botany 45: 755-756. Mulligan, G. A., and D. B. Munro. 1984. Chromosome numbers and sexual compatibility in North American Rorippa sylvestris (Cruciferae). Canadian Journal of Botany 62: 575-580. Mulligan, G. A., and D. B. Munro. 1990. Poisonous plants of Canada. Agriculture Canada Publication 1842/E. 96 pages. Mulligan, G. A., and A. E. Porsild. 1966. Rorippa calycina in the Northwest Territories. Canadian Journal of Botany 44: 1105-1106. Packer, J. G. 1964. Chromosome numbers and taxonom- ic notes on western Canadian arctic plants. Canadian Journal of Botany 42: 473-494. Pringle, J.S. 1969. Documented plant chromosome num- bers 1969:1. Sida 3: 350-351. Rollins, R.C. 1981. Weeds of the Cruciferae (Brassicaceae) in North America. Journal Arnold Arboretum 62: 517-540. Rollins, R. C. 1993. The Cruciferae of Continental North America. Stanford University Press, Stanford, California. 976 pages. Rousseau, C. 1968. Histoire, habitat et distribution de 220 plantes introduites au Québec. Le Naturaliste canadien 95: 49-169. Taylor, R. L., and G. A. Mulligan. 1968. Flora of the Queen Charlotte Islands, part 2, cytological aspects of the vascular plants. Queen’s Printer, Ottawa, Ontario. 148 pages. Warwick, S. L, L. D. Black, and J. K. Anderson. 1994. IOPB chromosome data 7. IOPB Newsletter 22: 4-5, Warwick, S. L., H. J. Beckie, A. G. Thomas, and T. McDonald. 2000. The biology of Canadian weeds, 8. Sinapis arvensis L. (updated), Canadian Journal of Plant Science 80; 939-961. Received 26 October 2001 Accepted 12 December 2002 T_T KT. 8 | « Additional Records of Bigeye Fishes (Priacanthidae) from the Atlantic Coast of Nova Scotia, Including the First Record of the Glasseye Snapper, Heteropriacanthus cruentatus BRIAN W. Coap! and JOHN GILHEN2 'Canadian Museum of Nature, P. O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 2Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Coad, Brian W., and John Gilhen. 2002. Additional records of bigeye fishes (Priacanthidae) from the Atlantic coast of Nova Scotia, including the first record of the Glasseye Snapper, Heteropriacanthus cruentatus. Canadian Field- Naturalist 116(4): 632-635. Three species of the bigeye fish family, Priacanthidae, are recorded from Nova Scotia based on new material. The speci- mens are all juveniles, one of the Glasseye Snapper, Heteropriacanthus cruentatus, eight of the Bigeye, Priacanthus are- natus, and two of the Short, Bigeye, Pristigenys alta. The record of the Glasseye Snapper is the first for Canada and a northward range extension of over 1000 km for the species. A key to the four species of western Atlantic bigeyes is given. Key Words: Glasseye Snapper, Heteropriacanthus cruentatus, first record, Nova Scotia, Canada, Bigeye, Priacanthus are- natus, Short Bigeye, Pristigenys alta, Priacanthidae The bigeyes (Priacanthidae) are a small family of carnivorous fishes, feeding on coral reef fishes, crus- taceans and worms. As their name indicates they have large eyes. The mouth is also large and very oblique with the lower jaw projecting. The scales are small and strongly ctenoid making bigeyes rough to the touch. Scales cover the body, head, the maxilla bone of the upper jaw and the lower jaw. The dorsal fin usually has 10 spines followed by 10-15 soft rays without a strong notch between spines and soft rays. The anal fin has 3 spines and 9 to 16 soft rays. The pelvic fin has one spine and five soft rays and is con- nected to the body by a membrane. There are 14 branched caudal fin rays. Bigeyes are usually bright red in colour (Coad et al. 1995). Bigeye fishes are usually nocturnal and are common on coral reefs and around rocky areas where they hide in crevices dur- ing the day. Adults may be found deeper than 400 metres. Young bigeyes are associated with floating debris near the surface in open ocean and are widely dispersed by ocean currents (Coad et al. 1995). Their occurrence along the coast of Nova Scotia is attribut- ed to transportation within eddies of Gulf Stream surface water. Like a number of juvenile tropical fishes known to occur during the summer months along the coast of Nova Scotia, they survive until autumn and die when water temperatures drop. The bigeye family of fishes is primarily circum- tropical in distribution. There are about 18 species known from the Atlantic, Indian and Pacific oceans. Only four species occur in the western Atlantic: Bulleye, Cookeolus japonicus (Cuvier, 1829), Big- eye, Priacanthus arenatus Cuvier, 1829, Short Bigeye, Pristigenys alta (Gill, 1862) and Glasseye Snapper, Heteropriacanthus cruentatus (Lacepéde, 1801). The first three species were reported from Nova Scotia by Scott and Scott (1988) and Coad et al. (1995). The latter species is reported for the first time from the Canadian Atlantic. Bigeye fishes are rarely recorded from the Atlantic coast of Nova Scotia. MacKay and Gilhen (1973) first recorded the capture of a 147.0 mm total length Bulleye [Cookeolus boops (Forster in Bloch and Schneider, 1801) = Cookeolus japonicus (Cuvier, 1829)] from Indian Point, St. Margarets Bay, Halifax County, on 16 September 1971 and a 41.7 mm approximate standard length Bigeye re- moved from the pharynx of a Pollock, Pollachius virens Linnaeus, 1758, caught on a hook and line at Gulch Shoal, off Hopson Island, Halifax County, on 8 September 1972. Scott and Scott (1988) included these and recorded a third species, a 22 mm total length Short Bigeye from near Corsair Canyon at 42°33'N, 64°35'W. In this paper, we record eight more specimens of Bigeye, two more specimens of Short Bigeye, and the first Canadian record of the Glasseye Snapper, all from the coastal waters of Nova Scotia in the 1990s. The record of the Glasseye Snapper confirms the presence of all four western Atlantic bigeye species in Canada. Catalogue Data Glasseye Snapper, Heteropriacanthus cruentatus (Figure 1) NSM (Nova Scotia Museum) /0037, 63.0 mm SL (standard length), from a mackerel trap, Red Bank, St. Margarets Bay, Halifax County, 44°37'32"N, 64°01'55”W, 18 October 1990, Eric Newton and Brian Coolen. This specimen is the first Canadian record. 632 2002 6cm FicureE 1. A juvenile (63.0 mm SL) Glasseye Snapper, Heteropriacanthus cruentatus, first record for the Canadian Atlantic, captured in a mackerel trap at Red Bank, St. Margarets Bay, Halifax County, Nova Scotia on 18 October 1990 by Eric Newton and Brian Coolen (NSM10037; negative N-24, 439). Bigeye, Priacanthus arenatus (Figure 2) NSM 10092, 70.6 mm SL, caught alive in a mackerel trap set off Red Bank, St. Margarets Bay, Halifax County, 44°37'32"N, 64°01'55”W, 17 September 1990, Eric Newton and Brian Coolen. This specimen is the second record for Nova Scotia. NSM 85350, 45.7 mm SL, found freshly dead at the strand line on the day after Hurricane Edward on Hirtle Beach, Lunenburg County, 44°27'05’N, 64°21'18"W, 3 September 1996, J. Sherman Bleakney. NSM 11796, 52.8 mm SL, NSM 11797, 50.2 mm SL and NSM 11798, 55.7 mm SL, caught alive mixed in with a large school of Butterfish, Peprilus COAD AND GILHEN: BIGEYE FISHES FROM NovA SCOTIA 633 triacanthus Peck, 1804, in a mackerel trap set off Joe Shatford’s Lobster Pound, Fox Point, St. Margarets Bay, Lunenburg County, 44°37'01"N, 64°03’00"”W, 10 October 1996, Eric Newton and Brian Coolen. NSM 85345, NSM 85346 and NSM 85347, 43.9-64.4 mm SL, washed up at high water mark, presumed to have been killed by a drop in sea temper- ature from 20°C to 14°C owing to a coastal upwelling occasioned by southwesterly winds, Clam Harbour Beach, Halifax County, 44°43’30”N, 62°53'25”W, 10 September 1999, Patrick Ryall. Short Bigeye, Pristigenys alta (Figure 3) NSM 10033, 19.4 mm SL, found freshly dead on sandy shore, Sambro Creek area, Indian Harbour, Halifax County, 44°28'17"N, 63°37'00”"W, 29 August 1990, Daniel Peters. This specimen is the second record for Nova Scotia NSM 10035, 25.3 mm SL, scooped up alive in a bucket of seawater, off Hopson Island, vicinity of Prospect, Halifax County, 44°27’00"N, 63°47’ 30" W, 31 August 1990, Ron Duggan. Descriptions of the Canadian Specimens Detailed accounts of the anatomy, colour and dis- tribution of these species may be found in Caldwell (1962) and Starnes (1988). All specimens are juve- niles and differ in colour from adults. Heteropriacanthus cruentatus The single specimen has a dorsal fin with rays X, 13 (10 spines and 13 soft rays), an anal fin with rays II, 14, a pectoral fin with 18 rays, 85 lateral series scales, 11 scales between the dorsal origin and the lateral line, and 23 total gill rakers. Head length (HL) in standard length (SL) is 2.8, head width in 6cm FicurE 2. A juvenile (70.6 mm SL) Bigeye, Priacanthus arenatus, second record for the Canadian Atlantic, captured in a mackerel trap at Red Bank, St. Margarets Bay, Halifax County, Nova Scotia on 17 September 1990 by Eric Newton and Brian Coolen (NSM 10092; negative N-24, 438) Figure 3. A juvenile (19.4 mm SL) Short Bigeye, Pristigenys alta, second record for the Canadian Atlantic, found freshly dead on sandy shore, Sambro Creek, Halifax County, 29 August 1990 by Daniel Peters (VSM /0033; negative N-25, 618). 634 HL is 1.9 (in SL 5.3), body width in SL is 5.6, head depth in SL is 2.5, body depth at the sixth dorsal fin spine level is 2.2 in SL, snout length in HL is 3.7 (10.3 in SL), orbital diameter in HL is 2.1 (6.0 in SL), interorbital distance in HL is 3.0 (8.3 in SL), length of the lower jaw in HL is 1.7 (4.7 in SL), pec- toral fin length in HL is 1.9 (5.3 in SL), pelvic fin length in HL is 1.5 (4.2 in SL), caudal peduncle length in SL is 6.9, caudal peduncle depth in SL 10.3, longest dorsal fin spine in SL is 6.1, longest dorsal fin soft ray in SL is 6.6, and longest anal fin soft ray in SL is 6.7. This specimen lacks scales on the chin and on the preopercle bone posterior to the shelf overlying the sensory canal, in contrast to the superficially similar Priacanthus arenatus. When freshly frozen it was a bright blue over the back and upper sides down to below the lateral line. The sides, from the back of the head to the base of the caudal fin, had six dark, uneven bands, interspersed with subtly paler elongate blotches which extend from the back to below the lat- eral line. The lower sides and belly were silvery-white with a distinct pinkish-red sheen. There were three small, dark-blue blotches on the lower sides above the anal fin. The eyes were a glossy orange-red. Fins were greyish at the base, with orange-brown spots, becom- ing blue-black along the outer edge, most prominently at the posterior edge of the caudal fin. In the preserved fish, the caudal fin shows rows of spots not seen in the Priacanthus arenatus juveniles. The dorsal and anal fins have a similar pigmentation to P. arenatus. The northernmost record of this species was a sin- gle record of a juvenile from off Cape May, New Jersey, itself a northward range extension from Southern Florida (Caldwell 1962; Starnes 1988). The Canadian record is a range extension of over 1000 km. Priacanthus arenatus The eight specimens have a dorsal fin with rays X, 14 (6 fish) or X, 15 (2), an anal fin with rays III, 15 THE CANADIAN FIELD-NATURALIST Vol. 116 - (6) or II, 16 (2) and a pectoral fin with 17 (3)or 18 (4) rays (one damaged). Total gill rakers are 29 (4) or 30 (4). Head length in SL is 2.8—2.9 (mean 2.9), pelvic fin length in HL is 1.1—1.4 (mean 1.2), and body depth at the sixth dorsal spine level is 2.4—2.7 (mean 2.5) in SL. NSM 10092 was a bright orange with a pinkish- purple sheen. The body and fins were peppered with small black dots which are probably due to a para- site. Pelvic fins were bordered in black and the pupil of the eye was deep blue. NSM 85350 was a pale pink with rosy bars. Preserved fish have the mem- branes of the spiny dorsal and anal fins darkly pig- mented while the anterior and basal parts of the soft dorsal and anal fins are less darkly pigmented. The pelvic soft rays and membranes are very darkly pig- mented while the spine is contrastingly clear. The body is brownish dorsally, silvery on the sides from scale reflections and silvery to whitish on the belly. The caudal fin may have a posterior margin band, particularly evident centrally. Pristigenys alta The two specimens have a dorsal fin with rays X, 11 (2), an anal fin with rays II, 10 (2), and a pec- toral fin with 17 (1) or 18 (1) rays. Total gill rakers are 26 (2). Head length in SL is 2.8—2.9 (mean 2.9), pelvic fin length in HL is 1.1—-1.2 (mean 1.2), and body depth at the fourth dorsal fin spine level is ban SEY There are no colour notes on fresh specimens of this species but the preserved material has a spiny dorsal fin with three bands which are partly reticu- late. The soft dorsal fin is darkened at its base. The anal fin spines and membranes are dark and the soft rays are dark basally. The pelvic fin spine has four bars and the soft rays and membranes are dark. The low counts of dorsal and anal soft fin rays and of lateral series scales are distinctive for this genus and species of bigeye. Key to the bigeyes (Priacanthidae) found in the Canadian Atlantic. 1A. Anal fin soft rays 9-11, usually 10; dorsal fin soft rays 10-12, usually 11; scales in lateral line series 45 or less — Pristigenys alta 1B. Anal fin soft rays 12-16; dorsal fin soft rays 12-15, usually 13-14; scales in lateral line series 58 or more — 2 2A. Pelvic fin length much greater than head length; scale rows between dorsal fin origin and lateral line 16-20; preorbital bone with strong anterior serrations — Cookeolus japonicus 2B. Pelvic fin length usually shorter than head length in young to slightly longer; scale rows between dorsal fin origin and lateral line less than 16; preorbital bone with fine anterior serrations — 3 3.A. Chin tip and posterior preopercle bone lacking scales; total gill rakers 21-25; anal fin soft rays 13-14, usually 14; dorsal fin soft rays 12-13, usually 13 — Heteropriacanthus cruentatus 3B. Chin tip and posterior preopercle bone scaled; total gill rakers 27-33; anal fin soft rays 14—16, usually 15; dorsal fin soft rays 13-15, usually 14 — Priacanthus arenatus 2002 Acknowledgments We are indebted to Wayne C. Starnes, Research Curator of Fishes, North Carolina State Museum of Natural Sciences, Raleigh, North Carolina, for his confirmation and correction of our tentative identifi- cations of these juvenile specimens. Eric Newton and Brian Coolen froze the specimens from their mackerel trap and donated them to the Nova Scotia Museum of Natural History for future study. Ron Duggan and J. Sherman Bleakney also kindly donat- ed their specimens. Patrick Ryall, Dalhousie Uni- versity, Halifax collected the Clam Harbour Beach specimens and Jamie Gibson and Ransom Myers, Dalhousie University, Halifax, Nova Scotia, donated this material to the Nova Scotia Museum of Natural History. The photographs were taken by Roger Lloyd, Learning Resources and Technology, Nova Scotia Department of Education, Halifax, Nova Scotia. COAD AND GILHEN: BIGEYE FISHES FROM Nova SCOTIA 635 Literature Cited Caldwell, D. K. 1962. Western Atlantic fishes of the fam- ily Priacanthidae. Copea 1962: 417-424. Coad, B. W., H. Waszezuk, and I. Labignan. 1995. Encyclopedia of Canadian Fishes. Canadian Museum of Nature, Ottawa, Ontario, and Canadian Sportfishing Productions, Waterdown, Ontario. viii + 929 pages. MacKay, K. T., and J. Gilhen. 1973. Hirundichthys ron- deleti, Cookeolus boops, Priacanthus arenatus, Seriola dumerili, four species new to the Canadian Atlantic ichthyofauna. Journal of the Fisheries Research Board of Canada 30 (12, part 1): 1911-1913. Scott, W. B., and M. G. Scott. 1988. Atlantic Fishes of Canada. Canadian Bulletin of Fisheries and Aquatic Sciences 219: xxx + 731 pages. Starnes, W. C. 1988. Revision, phylogeny and biogeo- graphic comments on the circumtropical marine percoid fish family Priacanthidae. Bulletin of Marine Science 43: 117-203. Received 12 April 2001 Accepted 12 December 2002 iP ee. SY - An Extraordinary Raccoon, Procyon lotor, Density at an Urban Park HENRY T. SMITH! and RICHARD M. ENGEMAN2 'Florida Department of Environmental Protection, Florida Park Service, 13798 S.E. Federal Highway, Hobe Sound, Florida 33455, USA 2National Wildlife Research Center, 4101 LaPorte Ave, Fort Collins, Colorado 80521-2154, USA Smith, Henry T., and Richard M. Engeman. 2002. An extraordinary Raccoon, Procyon lotor, density at an urban park. Canadian Field-Naturalist 116(4): 636-639. We report on a high density Raccoon (Procyon lotor) population at a 70.9 ha urban park in Fort Lauderdale, Florida, where in a 5-day period 160 raccoons were removed and 9 others observed without capture. This represents a minimum density of 238 raccoons/km?, and is one of the densest Raccoon populations reported. We discuss the ramifications of high density urban Raccoon populations relative to wildlife disease contagions and endangered species protection. Key Words: Raccoon Procyon lotor, endangered species protection, Florida, Fort Lauderdale, rabies, wildlife diseases. Hugh Taylor Birch State Park (HTBSP) is in Broward County, Florida, USA, within the limits of the City of Fort Lauderdale. It was initially obtained as public lands by the State of Florida on 31 December 1941. It consists of 56.7 ha of upland veg- etation, and 14.2 ha of freshwater and tidal wetlands, for a combined total of 70.9 ha. Terrestrial plant com- munities consist of disturbed maritime hammock and patchy areas dominated by exotic Australian Pine (Casuarina equisetifolia). HTBSP is completely con- tained within Fort Lauderdale which had a 1990 pop- ulation census of ca. 150 150 (Fort Lauderdale Community and Economic Development Depart- ment), and is surrounded by urban infrastructure to the north and south. The property is bordered to the east by additional infrastructure and the Atlantic Ocean; the Intracoastal Waterway (a large bulkhead- ed canal) truncates the entire western boundary. Raccoon (Procyon lotor) population densities in HTBSP have always been high since the first Southeast Florida district park biologist was hired in 1970 (R. Roberts, personal communication); howev- er, densities were not quantified. During the 1980s, tourists and local residents began the evening prac- tice of feeding Raccoons immediately outside the south boundary of the park, principally at the park drive entrance gate area along the Sunrise Boulevard right-of-way and the contiguous City Fire Station entrance. Although feeding of wildlife is unlawful inside all state parks, until November 2000 it was unregulated outside of HTBSP, including at the entrance. Efforts to control this practice included public environmental education about the detriments of feeding wildlife resources. Coarse, irregular esti- mates of the Raccoon population at the Park during the late 1990s ranged from 75 to 125 animals (H. Smith, personal observation). The high density Raccoon population presented a threat of epizootic disease transmission and bites to humans. High concentrations in urban areas and a willingness to accept food from humans by an ani- mal that many members of the public view as endearing create a much higher probability for human-animal contact than with many other wildlife species. The animals also presented constant vehicu- lar traffic problems in the evenings. A control pro- gram was conducted in November 2000 to reduce these hazards by reducing the abundance of Raccoons. Public sentiment restricted the approach to a “one-time only” live-trap and relocate program. This was cooperatively arranged between the Florida Department of Environmental Protection, Florida Fish and Wildlife Conservation Commission, and the City of Fort Lauderdale. Raccoons were captured using 80 X 25 X 30 cm Tomahawk live traps (P.O. Box 323, Tomahawk, Wisconsin 54487; use of prod- uct names does not constitute endorsement by the U.S. government, nor the Florida state government), baited with cat food. On the evening of 14 November, 123 Raccoons were captured and relocated by permit to a site south of Homestead, over 88 km away. On the evening of 15 November, HTBSP was again surveyed with 36 Raccoons observed. On the evening of 17 November, 37 additional Raccoons were trapped and relocated to the same site. On the evening of 18 November, active enforcement was initiated of City of Fort Lauderdale Ordinance No. C-00-48, Sec. 6-15 “Feeding of Wild Animals Prohibited.” A final survey that evening produced a count of nine raccoons near the park drive entrance gate. Thus, an absolute minimum number of Raccoons inhabiting the park area prior to removal efforts was 169. It is virtually impossible that any of the relocated Raccoons could have returned to be trapped more than once because of the great distance to the reloca- tion site, the habitat, including waterways and bridges they would have to cross through urban Fort 636 2002 Lauderdale, and the short (5 day) time frame in which all work was completed. Thus, the minimal Raccoon density for HTBSP at the time this work was implemented was 238/km?. Artificially enhanced populations of Raccoons are a problem in wildlands and urban-suburban areas of Florida (Heugel 1991*), and on a more cosmopolitan level (e.g., Rosatte et al. 1992a). Due to the high degree of habitat alteration, fewer species inhabit urban environments than natural environments. However, for those species that inhabit urban envi- ronments, increased availability and concentration of food, den sites or other refuges may induce dense populations (e.g., Dickman 1987; Dickman and Doncaster 1987; Riley et al. 1998). The highest Raccoon density we could find in the literature came from one small (30 ha) section in Rock Creek Park (710 ha total) in Washington, D.C., where the density was estimated as high as 333/km’, and estimated Raccoon densities from other study sites in the same park ranged as high as 172/km? (Riley et al. 1998). Estimated densities of raccoons in metropolitan Toronto, Ontario, ranged as high as 140/km? (Rosatte et al. 1992a). The minimal poten- tial density we observed for HTBSP ranks as one of the densest congregations of Raccoons recorded (see Riley et al. 1998 for a range of Raccoon densities). Its Raccoon density was over 4 to 200 times as great as reported rural Raccoon densities (0.9-55.6/km7) (Mech et al. 1968; Johnson 1970; Urban 1970; VanDruff 1971; Sonenshine and Winslow 1972; Lehman 1977*; Keeler 1978; Lehman 1980*; Orloff 1980*; Slate 1980*; Rabinowitz 1981; Jacobson 1982; Nottingham et al. 1982; Lehman 1984%*; Moore and Kennedy 1985; Leberg and Kennedy 1988; Perry et al. 1989; Hasbrouck 1991; Kennedy et al. 1991; Hable et al. 1992; Nixon et al. 1993*). The high rural density of 250/km? reported by Twichell and Dill (1949) may be an anomaly for rural populations as it too was 4-200 times greater than the other reported rural densities at 0.9- 55.6/km2. Rosatte et al. (1992b) and Rosatte (2000) demonstrated the patchiness of Raccoon densities in urban habitats when large areas are trapped. Many of the high densities reported, including our observa- tions, involved small urban parks presenting large numbers of Raccoons. Densities outside HTBSP were probably much lower. High densities of Raccoons at HTBSP posed a concern for various reasons. First, a number of dis- eases are endemic in raccoon populations, in particu- lar, rabies (Winkler and Jenkins 1991). Raccoons are the most common vectors of rabies in the U.S. (Krebs et al. 2001), and two-thirds of all rabid ani- mals in Florida have been Raccoons (Burridge et al. 1986). A large number of uncommonly reported species have been found rabid in association with Raccoon rabies epizootics (Burridge et al. 1986), in SMITH AND ENGEMAN: EXTRAORDINARY RACCOON DENSITY 637 part due to Raccoon usage of urban resources (Winkler and Jenkins 1991). Urban Raccoons are of particular concern for wildlife disease control (e.g., Broadfoot et al. 2001). Most human exposures have been largely due to failure to apply common sense in interacting with Raccoons (Jenkins and Winkler 1987; Jenkins et al. 1988). It follows that a high-den- sity Raccoon population in an urban setting, espe- cially where people have become enamored of them or accustomed to feeding them, as at HTBSP, pre- sents a high-risk recipe for human exposure to rabies during an outbreak (e.g., Kappus et al. 1970; Jenkins and Winkler 1987; Jenkins et al. 1988). We would not expect future high-density urban Raccoon situa- tions in Florida to be addressed with a trap and relo- cate program due to legal and wildlife management issues. However, if another exception is made, all animals should be vaccinated for rabies prior to release (Rosatte et al. 1992b); otherwise there is a danger of artificially spreading rabies as has been reported previously (Jenkins and Winkler 1987). Ear-tagging would help monitor their movements post-release. In addition, Raccoons in Florida have become an example of an abundant native vertebrate that impacts the conservation of endangered species (e.g., Garrott et al. 1993) as they cause substantial destruc- tion of sea turtle nests in Florida and throughout the southeastern United States (Stancyk 1982; Engeman et al. in press), as well as to shore-nesting birds such as terns and skimmers (Rodgers et al. 1996; Thompson et al. 1997). High density urban Raccoon populations in close proximity to beaches where nesting takes place may provide a reservoir of ani- mals that artificially intensifies predation (Engeman et al. in press), and requires predator management to reduce their negative impacts (e.g., Reynolds and Tapper 1996; Hecht and Nickerson 1999). Acknowledgments We thank T. Hardwick for designing and imple- menting the trapping program under contract to FDEP. J. Higgins of FDEP arranged the trapping contract and ensured other aspects ran smoothly. T. Regan of the Florida Fish and Wildlife Conservation Frosbutter of FDEP helped inform the City of Ft. Lauderdale Commissioners on the detrimental effects of feeding wildlife resources. G. Jones of FDEP supported the project, despite public contro- versy. F. Cook, K. Fagerstone, M. Pipas, and R. Rosatte provided valuable reviews of the manuscript. Documents Cited (marked * in text citation) Huegel, C. N. 1991. Raccoons. Report SS-WIS-34, Florida Cooperative Extension Service, University of Florida, Largo, Florida. Lehman, L. E. 1977. Population ecology of the raccoon of the Jasper-Pulaski wildlife study area, Pittman- 638 Robertson Bulletin 9, Indiana Department of Natural Resources, Indianapolis, Indiana. Lehman, L. E. 1980. Raccoon population ecology on the Brownstown Ranger District, Hoosier National Forest. Pittman-Robertson Bulletin 11, Indiana Department of Natural Resources, Indianapolis, Indiana. Lehman, L. E. 1984. Raccoon density, home range, and habitat use on south-central Indiana farmland. Pittman- Robertson Bulletin 15, Indiana Department of Natural Resources, Indianapolis, Indiana. Nixon, C. M., J. B. Sullivan, R. Koerkenmeier, T. Esker, G. R. Lang, L. L. Hungerford, M. Mitchell, G. A. Demonceaux, G. F. Hubert, Jr., and R. D. Bluett. 1993. Illinois raccoon investigations. Final Report, Project Number W-104-R- 1,2,3, Center for Wildlife Ecology, Illinois Natural History Survey, Champaign, Illinois. Orloff, S. 1980. Raccoon status in Contra Costa and Alameda counties. Job Progress Report, Project Number W-54-R-12, California Department of Fish and Game, Nongame Wildlife Investigations, Sacramento, California. Slate, D. 1980. Raccoon population dynamics in different habitat types in New Jersey. Fish and Wildlife Reference Service Document MIN 298280351, U.S. Fish and Wildlife Service, Bethesda, Maryland. Literature Cited Broadfoot, J. D., R. C. Rosatte, and D. T. O’Leary. 2001. Raccoon and skunk population models for urban disease control planning in Ontario, Canada. Ecological Applications 11: 295-303. Burridge, M. J., L. A. Sawyer, and W. J. Bigler. 1986. Rabies in Florida. Department of Health and Rehabili- tative Services, Tallahassee, Florida. 19 pages. Dickman, C. R. 1987. Habitat fragmentation and verte- brate species richness in an urban environment. Journal of Applied Ecology 24: 337-351. Dickman, C. R., and C. P. Doncaster. 1987. The ecology of small mammals in urban habitats. I. Populations in patchy environments. Journal of Animal Ecology 56: 629-640. Engeman, R., B. Constantin, R. Noel, and J. Woolard. In press. Monitoring predators to optimize turtle nest protection through control. Biological Conservation. Garrott, R. A., P. J. White, and C. A. White. 1993. Overabundance: an issue for conservation biologists? Conservation Biology 7: 946-949. Hable, C. P., A. N. Hamir, D. E. Snyder, R. Joyner, J. French, V. Nettles, C. Hanlon, and C. E. Rupprecht. 1992. Prerequisites for oral immunization of free-rang- ing raccoons (Procyon lotor) with a recombinant rabies virus vaccine: study site ecology and bait system devel- opment. Journal of Wildlife Disease 28: 64-79. Hasbrouck, J. J. 1991. Demographic responses of rac- coons to varying exploitation rates. Ph.D. thesis, Iowa State University, Ames Iowa. Hecht, A., and P. R. Nickerson. 1999. The need for preda- tor management in conservation of some vulnerable species. Endangered Species Update 16: 114-118. Jacobson, J. E. 1982. Parasite relationships between an urban and a rural raccoon population. M.S. thesis. Purdue University, West Lafayette, Indiana. Jenkins, S. R., and W. G. Winkler. 1987. Descriptive epidemiology from an epizootic of raccoon rabies in the THE CANADIAN FIELD-NATURALIST RIEL LL Vol. 116 - Middle Atlantic States, 1982-1983. American Journal of Epidemiology 126: 429-437. Jenkins, S. R., B. D. Perry, and W. G. Winkler. 1988. Ecology and epidemiology of raccoon rabies. Review Infectious Diseases 10: S620—S625. Johnson, A. S. 1970. Biology of the raccoon in Alabama. Auburn University Experiment Station Bulletin 402. Kappus, K. D., W. J. Bigler, R. G. McLean, and H. A. Trevino. 1970. The raccoon as an emerging rabies host. Journal of Wildlife Diseases 6: 507-509. Keeler, E. W. 1978. Some aspects of the natural history of the raccoon (Procyon lotor) in Cades Cove, The Great Smokies National Park. M.S. thesis. University of Tennessee, Knoxville, Tennessee. Kennedy, M.L., J. P. Nelson, Jr., F. W. Weckerly, D. W. Sugg, and J. C. Stroh. 1991. An assessment of selected forest factors and lake level in raccoon manage- ment. Wildlife Society Bulletin 19: 151-154. Krebs, J. W., A. M. Mondul, C. E. Ruprecht, and J. E. Childs. 2001. Rabies surveillance in the United States during 2000. Journal of the American Veterinary Medicine Association 219: 1687-1699. Leberg, P. L., and M. L. Kennedy. 1988. Demography and habitat relationships of raccoons in western Tennessee. Proceedings of the Annual Conference of the Southeastern Association of Fish and Wildlife Agencies 42: 272-282. Mech, L. D., D. M. Barnes, and J. R. Tester. 1968. Seasonal weight changes, mortality, and population structure of raccoons in Minnesota. Journal of Mammalogy 49: 63-73. Moore, D. W., and M. L. Kennedy. 1985. Factors affecting response of raccoons to traps and population size estimation. American Midland Naturalist 114: 192-197. Nottingham, B. G., K. G. Johnson, J. W. Woods, and M. R. Pelton. 1982. Population characteristics and harvest relationships of a raccoon population in east Tennessee. Proceedings of the Annual Conference of the Southeastern Association of Fish and Wildlife Agencies 36: 691-700. Perry, B. D., N. Garner, S. R. Jenkins, K. McCloskey, and D. H. Johnston. 1989. A study of techniques for distribution of oral rabies vaccine to wild raccoon popu- lations. Journal of Wildlife Diseases 25: 206-217. Rabinowitz, A. R. 1981. The ecology of the raccoon (Procyon lotor) in Cades Cove, The Great Smokies National Park. Ph.D. thesis. University of Tennessee, Knoxville, Tennessee. Reynolds, J. C., and S.C. Tapper. 1996. Control of mammalian predators in game management and conser- vation. Mammal Review 26: 127-156. Riley, S. P. D., J. Hadidian, and D. A. Manski. 1998. Population density, survival, and rabies in raccoons in an urban national park. Canadian Journal of Zoology 76: 1153-1164. Rodgers, J. A., Jr., H. W. Kale, II, and H. T. Smith. Editors. 1996. Rare and endangered biota of Florida, Volume V: Birds. University Press of Florida, Gaines- ville, Florida. 688 pages. Rosatte, R. 2000. Management of raccoons (Procyon lotor) in Ontario, Canada: Do human intervention and disease have significant impact on raccoon populations? Mammalia 64: 369-390. Rosatte, R., M. J. Power, and C. D. MacInnes. 1992a. 2002 Density, dispersion and habitat of skunks (Mephitis mephitis) and raccoons (Procyon lotor) in metropolitan Toronto. Pages 932-944 in Wildlife 2001: Populations. Edited by D. R. McCullogh and R. H. Barret. Elsevier Applied Science, London. 1163 pages. - Rosatte, R. C., M. J. Power, C. D. MacInnes and J. B. Campbell. 1992b. Trap-vaccinate-release and oral vac- cination for rabies control in urban skunks, raccoons and foxes. Journal of Wildlife Diseases 28: 562-571. Sonenshine, D. E., and E. L. Winslow. 1972. Contrasts in distribution of raccoons in 2 Virginia localities. Jour- nal of Wildlife Management 36: 838-847. Stancyk, S. E. 1982. Non-human predators of sea turtles and their control. Pages 139-152 in Biology and Conservation of Sea Turtles. Edited by K. A. Bjorndal. Smithsonian Institution Press, Washington, D.C. Thompson, B. C., J. A. Jackson, J. Burger, L. A. Hill, E. M. Kirsch, and J. L. Atwood. 1997. Least Tern (Sterna antillarum). Birds of North America account 290. Edited by A. Poole and F. Gill. The Academy of SMITH AND ENGEMAN: EXTRAORDINARY RACCOON DENSITY 639 Natural Sciences, Philadelphia, Pennsylvania, and The American Ornithologist’s Union, Washington, D.C. 32 pages. Twichell, A. R., and H. H. Dill. 1949. One hundred rac- coons from one hundred and two acres. Journal of Mam- malogy 30: 130-133. Urban, D. 1970. Raccoon populations, movement pat- terns, and predation on a managed waterfowl marsh. Journal of Wildlife Management 34: 372-382. VanDruff, L. W. 1971. The ecology of the raccoon and opossum, with emphasis on their role as waterfowl nest predators. Ph.D. thesis, Cornell University, Ithaca, New York. | Winkler, W. G., and S. R. Jenkins. 1991. Raccoon rabies. Pages 325-340 in The Natural History of Rabies, 2™4 Edition. Edited by G. M. Baer. CRC Press, Boca Raton, Florida Received 10 August 2001 Accepted 14 January 2003 a Notes Initial Beneficial Effects of Fire and Bulldozing on Neglected Milkvetch, Astragalus neglectus, in an Eastern Ontario Alvar Woodland PAUL M. CATLING and ADRIANNE SINCLAIR Biology Department, University of Ottawa, 30 Marie Curie Street, P.O. Box 450, Station A, Ottawa, Ontario KIN 6N5 Catling, Paul M., and Adrianne Sinclair. 2002. Initial beneficial effects of fire and bulldozing on Neglected Milkvetch, Astragalus neglectus, in an eastern Ontario alvar woodland. Canadian Field-Naturalist 116(4): 640-642. Following a fire 16 months earlier, data on presence and amount of a rare and restricted plant, Astragalus neglectus, was collected in order to determine the initial effects of burning and bulldozing. Frequency and cover were recorded in 340 one metre square quadrats of which 150 were in burned woodland, 90 were along the undriven portions of two bulldozed tracks through unburned woodland, and 100 were in unburned woodland. In addition, the number of Astragalus neglectus seedlings was recorded within 2.5 m of each of 450 observation points, each 5 m apart, including 150 along two bulldozed tracks and 300 in adjacent woodland. There were significantly more occurrences of Astragalus neglectus in burned wood- lands and bulldozed tracks than in adjacent unburned woodlands. More than 95% of the Astragalus neglectus plants record- ed were definitely seedlings, and the percentage seedlings may have been as high as 99%. A major spread and population increase in Astragalus neglectus is therefore associated with the recent disturbance events. This provides additional accu- mulating evidence that disturbance management may be important in maintaining high native biodiversity and rare species in alvar landscapes. Key Words: alvar, limestone, fire, flora, biodiversity, diversity, rare species, succession, management, biomass removal, burn, burning, bulldozing, Great Lakes, Ontario, Canada Neglected Milkvetch (Astragalus neglectus (Torr. & A. Gray) E. Sheld.) is considered rare to uncom- mon in Ontario (S3, Newmaster et al. 1998) and is a rare plant locally and regionally in the Ottawa valley where it has a relatively large number of occurrences (Brunton 1986*; Brownell and Larson 1995*, Brunton 1997*). Over 70% of the Ontario locations for this plant are in alvar landscapes (Catling and Brownell 1995; Brownell and Riley 2000) where the plants grow in open or semi-open locations on thin soil over limestone rock (Catling 1995). A year after a fire on the Burnt Lands alvar northwest of Ottawa in eastern Ontario, it was noticed that burned alvar woodland and tracks bulldozed through unburned alvar woodland contained large numbers of young plants of Neglected Milkvetch. This observation pro- filed questions regarding the use of disturbance in management of alvars and protection of distinctive alvar flora. Many plant communities of alvar landscapes are currently recognized as globally imperiled and alvars contain numerous rare species (Catling and Brownell 1999; Brownell and Riley 2000), but information to effectively manage these communities is currently inadequate. A few recent studies however have sug- *see Documents Cited gested that large scale biomass removal may signifi- cantly contribute to biodiversity protection (Catling and Brownell 1998; Catling et al. 2001, 2002). The objective of the work reported here was to determine the extent to which fire and bulldozing had an initial beneficial effect on Neglected Milkvetch in order to contribute to understanding of the effects of distur- bance and develop appropriate management plans to maximize protection of biodiversity in alvar land- scapes. Methods Study area The Burnt Lands alvar is located in the county of Lanark and the former Regional Municipality of Ottawa-Carleton (see Brownell and Riley 2000), now part of city of Ottawa, on the Smith’s Falls Limestone Plain in eastern Ontario. The sites specifi- cally studied are centred on 45.2559 °N, 76.1432°W, and detailed descriptions of the actual positioning of transects are on file with the Kemptville office of the Ontario Ministry of Natural Resources. On 23 June 1999, 152 ha of alvar woodland in the Burnt Lands was burned during a major fire. Dry conditions in the relatively shallow soil over porous limestone rock probably led to a relatively slow return to a mixed boreal and fire prone forest type 640 2002 following the last fire in 1870. It seems likely that fire and post-fire succession has been going on in the area for many hundreds (or thousands) of years. The study area (White 1979; Brunton 1986*; Brownell and Riley 2000) included burned and unburned alvar woodland dominated by Northern White Cedar (Thuja occidentalis L.), White Spruce (Picea glauca (Moench) Voss), Balsam Fir (Abies balsamea (L.) Miller), White Pine (Pinus strobus L.), and Quaking Aspen (Populus tremuloides Michx.). In order to contain the fire, accessibility was improved by bulldozing tracks which resulted in scraping of the shallow, often gravelly soil to 1 dm deep, with organic material, into piles sometimes with woody debris including broken trees and branches. Where the disturbance to soil as a result of scraping was minimal there was still compaction and removal of all woody plants. In general, the flatness made development of a system of tracks for fire- fighting very easy. Since the plants were without flowers we used leaf pubescence to distinguish species. Astragalus neglectus has leaves that are glabrous above while A. canadensis has leaves that are hairy above. This dif- ference appears to be a satisfactory discriminator based on examination of 30 correctly identified flowering specimens of each species collected throughout Ontario (in the Agriculture and Agri-food Canada herbarium (DAO) in Ottawa). Although A. canadensis was abundant along one track, it was confined to an area of approximately 100 m? and was not found in either the burned or unburned woodland. Sampling procedures The sampling was done in October 2000. The sampling time greatly facilitated the observation of Astragalus spp. because they remained green after the first few autumn frosts when many other species that they occurred with (Aster macrophyllus etc.) had turned brown. The sampling procedures involved (1) quadrats and (2) observation points: (1) 340 one-metre-square quadrats, 5 m apart, were laid out along transects in three examples of burned woodland (50 quadrats each), along the undriven portions of two bulldozed tracks (45 quadrats each), and in examples of two unburned woodland (50 quadrats each). The sampled unburned woodland was 10 to 500 m from the burned and bull- dozed sampling sites and at similar elevation. It was evidently of the same composition (see above) based on the identity of the burned or bulldozed trees. Data included presence and cover of Astragalus neglectus in all quadrats. (2) the number of seedling Astragalus neglectus was recorded within 2.5 m of each of 450 observation points each 5 m apart. Of these observation points, 150 were along two bulldozed tracks and 300 were along transects in the adjacent woodland on either side of NOTES 641 these bulldozed tracks, but 10 m from the edge. This latter distance ensured that the effect of bulldozing was not a factor relating to occurrence in the woods, but was near enough to the track to minimize the effect of other factors such as soil depth and moisture content. Yates’ corrected Chi-square was used to evaluate significance since 2 X 2 tables were used. Mean fre- quencies provided expected values. Results More than 95% of the Astragalus neglectus plants recorded were definitely derived from seed that ger- minated earlier in the year. Frequency of occurrence of Astragalus neglectus plants in burned woods was 25.33% (38) and along bulldozed tracks was 17.7% (16), compared to 1.3% (2) in unburned woodland. These differences are significant (Chi-square 0.001, 0.009 respectively). Cover in quadrats in burned woodland was 145 dm’, along bulldozed tracks 49.5 dm? and was 0.5 dm? in unburned woodland. Adjustments of these total cover values for 100 quadrats are 96.6, 55.0 and 0.5 dm. Astragalus neglectus was recorded at 41.66 % (61) of observation points along bulldozed tracks and at 1.66% (5) of observation points in unburned woodland. These differences are significant below the 0.0001 level. Only seven Astragalus neglectus seedlings were recorded in the unburned woods, but 1177 seedlings were recorded at the 61 observation points along bulldozed tracks. Discussion The significantly higher occurrences of Astragalus neglectus in burned woodlands and bulldozed tracks than in adjacent unburned woodlands suggest an ini- tial beneficial effect of burning and bulldozing. The high percentage of this perennial plant, which was a result of seedlings in newly created habitat, indicates a major spread and population increase in Astragalus neglectus following the recent disturbance events. This spread and increase may be promoted by reduced competition for seedlings and/or increased nutrients leading to increased fecundity of survivors and increased survival of seedlings. In addition distur- bance may be necessary to promote seed germination. Astragalus neglectus was highlighted as one of a number of rare species occupying a floristically rich area of the Burnt Lands that had been burned 37 years earlier (Catling and Brownell 1998). The data presented provide more substantial evidence for the importance of disturbances such as fire and bulldoz- ing in alvar landscapes, and the importance of transi- tional disturbed plant communities such as succes- sional alvar burns. Maintenance of both maximum native biodiversity and rare species like Astragalus neglectus on alvars will probably require disturbance management techniques. Only recently has the potential dependence on disturbance by a portion of 642 the alvar flora and fauna been adequately recog- nized. However, parts of all Ontario alvars studied in depth have been burned, and burning is a natural process, so these conclusions are not surprising. Acknowledgments Funding for this study was provided by the Southcentral Sciences Section of the Ontario Ministry of Natural Resources (OMNR). Aerial pho- tography was supplied by OMNR, and field equip- ment, other supplies and resources and management help was provided by the University of Ottawa. Don Cuddy of the Ontario Ministry of Natural Resources provided much valuable support. Documents Cited (marked * in the text) Brownell, V. R., and B. M. H. Larson. 1995. An evalua- tion framework for natural areas in the regional munici- pality of Ottawa-Carleton. Volume 2, technical appen- dices. Regional Municipality of Ottawa-Carleton, Planning and Property Department 28-07. 91 pages. Brunton, D. F. 1986. A life science inventory of the Burnt Lands. Ontario Ministry of Natural Resources. 118 pages + maps. Brunton, D. F. 1997. Distributionally significant vascular flora of the regional municipality of Ottawa-Carleton, Ontario. 31 pages. Literature Cited Brownell, V. R., and J. L. Riley. 2000. Alvars of Ontario: significant alvar natural areas in the Ontario Great Lakes region. Federation of Ontario Naturalists, Toronto. 269 pages. THE CANADIAN FIELD-NATURALIST [EL .TTt«* Vol. 116 - Catling, P. M. 1995. The extent of confinement of vascu- lar plants to alvars in southern Ontario. Canadian. Field- Naturalist 109: 172-181. Catling, P. M., and V. R. Brownell. 1995. A review of the alvars of the Great Lakes region: distribution, floris- tic composition, phytogeography and protection. Canadian. Field-Naturalist 109: 143-171. Catling, P. M., and V. R. Brownell. 1998. Importance of fire in alvar ecosystems — evidence from the Burnt Lands, Eastern Ontario. Canadian Field-Naturalist 112: 661-667. Catling, P. M., and V. R. Brownell. 1999. The alvars of the Great Lakes region. Pages 375-391 in Savanna, bar- ren and rock outcrop communities of North America. Edited by R. C. Anderson, J. S. Fralish, and J. M. Baskin. Cambridge University Press. Cambridge, United Kingdom. Catling, P. M., A. Sinclair, and D. Cuddy. 2001. Vascular plants of a successional alvar burn 100 days after a severe fire and their mechanisms of re-establish- ment. Canadian Field-Naturalist 115: 214-222. Catling, P. M., A. Sinclair, and D. Cuddy. 2002. Plant community composition and relationships of disturbed and undisturbed alvar woodland. Canadian Field- Naturalist 116: 571-579. Newmaster, S. G., A. Lehela, P. W. C. Uhlig, S. MeMurray, and M. J. Oldham. 1998. Ontario Plant List. Forest Research Information Paper 123. Ontario Ministry of Natural Resources, Peterborough, Ontario. White, D. J. 1979. Burnt Lands alvar. Trail and Landscape 13: 34-38. Received 6 February 2001 Accepted 19 December 2002 Successful Spawning by Chinook Salmon, Oncorhynchus tshawytscha, in the St. Lawrence River at Cornwall, Ontario M. BRIAN C. HICKEY St. Lawrence River Institute of Environmental Sciences, Windmill Point, 2 Belmont Street, Cornwall, Ontario K6H 4Z1 Canada Hickey, M. Brian C. 2002. Successful spawning by Chinook Salmon, Oncorhynchus tshawytscha, in the St. Lawrence River at Cornwall, Ontario. Canadian Field-Naturalist 116(4): 642-645. Between 26 October and 3 November, 2000, I observed Chinook Salmon (Oncorhynchus tshawytscha) spawning in a human-made creek entering the St. Lawrence River at Cornwall, Ontario. During the following spring and summer I searched for signs of successful spawning by conducting visual searches supplemented with regular seine netting. I cap- tured young-of-the-year Chinook Salmon on four separate occasions, between June and September 2001. This is the second published report of successful spawning by Chinook Salmon in the St. Lawrence River and the first to identify a successful spawning location. Key Words: Chinook Salmon, Oncorhynchus tshawytscha, spawning, St. Lawrence River, Cornwall. Chinook Salmon (Oncorhynchus tshawytscha) are native to the west coast of North America (Hart 1973) but have been introduced to the Great Lakes on several occasions, dating back to the late 1800s (MacKay 1969; Scott and Crossman 1973). These repeated introductions have failed to produce a self- sustaining Chinook population in the Great Lakes, and although Chinook are now widespread through- 2002 out the Great Lakes-St. Lawrence River, they require continued stocking to maintain a sport fishery (MacKay 1969; Scott and Crossman 1973; Hubbs and Lagler 1974). In the St. Lawrence River, Chinook Salmon are occasionally captured by anglers, but there are no known spawning locations on the Ontario side of the river. Ribey and Chapleau (1996) captured young- of-the year Chinook Salmon while seining in the lit- toral zone of the St. Lawrence River near Cornwall, Ontario. Based on the small size (36.8 and 42 mm) of the two individuals they captured, they concluded that the fish represented local spawning by Chinook Salmon (Ribey and Chapleau 1996). Here, I extend the observation of Ribey and Chapleau (1996) by identifying a Chinook spawning site in the St. Lawrence River at Cornwall. My observations include records of fall spawning behaviour by pairs of adult Chinook Salmon, capture of gravid females, and the presence of young during the following spring, summer and fall. On three separate occasions, between 26 October 2000 and 3 November 2000, I observed adult Chinook Salmon in a human-made creek near down- town Cornwall, Ontario (45°00'45"N; 74°44'00”W). oF gS haga rg ter ce ra NOTES 643 The 200 m long creek originates at the old Cornwall Canal and meanders though the Rotary Eco Garden for Conservation and Recreation before entering the St. Lawrence River. The creek and the surrounding naturalized areas were constructed, in 1997, during the implementation of the Remedial Action Plan for the St. Lawrence River Area of Concern (Dreier et al. 1997). The creek (Figure 1) varies in depth from about 0.2 to 0.8 m, and in width from about 4 to 6 m. At the upstream end of the creek there is a small waterfall that prevents the further upstream migration of fish into the Cornwall canal. The substratum of rounded cobble and gravel, and the meander radii were designed to mimic a natural stream with alternat- ing shallow riffle areas and deeper pools. The water in the canal, which serves as the source of the water for the creek, originates as a small diversion from the St. Lawrence River near the Moses- Saunders power dam and therefore water quality in the creek is similar to that of the St. Lawrence River. Mean (+ s.d.) monthly temperatures °C in the creek were: April, 6.8 + 2.1(n = 6); May, 12.8 (n= 1); June, 18.0+ 4.6 (n =3); July, 22.0 + 0.9 (n = 3), August, 23.7 (n= 1). The adult salmon I observed were identified as Chinook based on their large size and the spotted —— a 2 ay ir rr ss -PR RP OU ERC EP GUE ‘ of J y ! eaillll OUP a} LD a Hn! FiGuRE 1. Human-made creek at the Rotary Eco Garden for Conservation and Recreation in Cornwall, Ontario, Spawning behaviour of Chinook Salmon was first observed in October 2000 in the area under the bridge shown in the photograph 644 pattern on the dorsal and caudal fins (Scott and Crossman 1973; Hart 1973). One individual, which I captured with a dip net, measured approximately 90 cm in total length. On two visits to the creek I observed pairs of Chinook engaged in spawning activity (excavating nests, swimming side-by-side), and one dead female Chinook with eggs (presumably killed by vandals) found by a City of Cornwall staff member. To determine whether spawning was successful, I conducted regular searches (1—6 searches per month) of the creek between May and September 2001. These visual searches were supplemented with seine netting once or twice per month. On four separate occasions I captured young-of- the-year Chinook Salmon. On 14 June 2001, I cap- tured a single Chinook. On 27 June 2001, I captured a second individual which was preserved as a vouch- er specimen and deposited in the collection of the Canadian Museum of Nature (catalogue number CMNFI 2001-0055). On 16 July 2001, I captured three specimens, and on 25 September 2001, I cap- tured an additional 11 Chinook Salmon. Specimens were identified using the keys and descriptions in Scott and Crossman (1973) and Hart (1973). The identity of the voucher specimen was confirmed by Sylvie Laframboise and Claude Renaud of the Canadian Museum of Nature (Ichthyology). The pigmentation around the edge of the adipose fin and 10 long broad parr marks were distinctive of young Chinook Salmon. The total length of the voucher specimen was 57 mm. To minimize the stress to the fish I returned the salmon to the water as quickly as possible and did not take measurements of individual fish; however, the 11 fish I captured on 25 September 2001 were approximately 100-150 mm in total length (Figure 2). FIGURE 2. One of 11 young-of-the-year Chinook Salmon captured on 25 September 2001 in the creek at the Rotary Eco Garden for Conservation and Recreation in Cornwall, Ontario. THE CANADIAN FIELD-NATURALIST Vol. 116 - This is the first documentation of a location of successful spawning by Chinook Salmon in the St. Lawrence River basin. The precise origin of the young-of-the-year Chinook Salmon reported by Ribey and Chapleau (1996) is unknown, but it could not have been the creek that I describe here, since their observations were made before the Rotary Eco Garden and creek were constructed. Chinook Salmon runs occur in several of the tributaries on the U.S. side of the St. Lawrence but their success has not been documented (S. Lepan; New York Department of Environmental Conservation, Cape Vincent Fisheries Station, personal communication). The results I report here, along with those of Ribey and Chapleau (1996), suggest that natural reproduc- tion by Chinook Salmon in the St. Lawrence River may be more widespread than previously thought. Whether the spawning frequency and success rates are sufficient to result in a self-sustaining population of Chinook requires more detailed information. Acknowledgments I am grateful to all those who participated in the Rotary Eco Garden habitat project, especially Norm Levac and John de Ronde (City of Cornwall) and Mike Eckersley (Ontario Ministry of Natural Resources) who undertook most of the design work. The project was constructed with funding from the Great Lakes 2000 Cleanup Fund (Environment Canada), the City of Cornwall, the Cornwall Rotary Club, Canada Trust Friends of the Environment Foundation and John and Valery Markell. Andy Code (Raisin Region Conservation Authority) provided encouragement and logistical support during the construction of the creek. Jamie Davidson, Luc Landry-Deptuck, Jeff Ridal, Heather Moore, Katimavik participants (Jeff Chik, Karena O’Brien, and Marie-Josée Comeau) and participants of the Rotary Adventures in the Environment (Jodie Lalonde, Sarah Tucker, Jessica Davey-Quastick, Shelagh McGuinty, Sean Dunn, Lindsey Finnerty, Tim VanWinckle, Seamus McGrath) assisted with the fish surveys. I thank Heather Moore, Jeff Ridal, Jonathan R. Hill, Rob Gratton, Jennifer Gibson and an anonymous reviewer for providing constructive com- ments on the manuscript. Literature Cited Dreier, S.I., J. Anderson, J. Biberhofer, M. Eckersley, R. Helliar, M. B. C. Hickey, L. Richman, F. Stride and The St. Lawrence River Remedial Action Plan Public Advisory Committee. 1997. Great Lakes Great River: Remedial Action Plasn for the St. Lawrence River (Cornwall) Area of Concern. 203 pages. ’ Hart, J. L. 1973. Pacific Fishes of Canada. Bulletin 180. Fisheries Research Board of Canada. Ottawa. 740 Pages. Hubbs, C. L., and K. F. Lagler. 1974. Fishes of the Great Lakes Region. University of Michigan Press. Ann Arbor. 213 pages. 2002 NOTES 645 MacKay, H. H. 1969. Fishes of Ontario. The Ontario Department of Lands and Forests. 292 pages. Ribey, S. R., and F. Chapleau. 1996. Evidence of suc- cessful Chinook Salmon, Oncorhynchus tshawytscha, spawning in the St. Lawrence River, near Cornwall, Ontario. Canadian Field-Naturalist 110: 346-347. Scott, W. B., and E. J. Crossman. 1973. Freshwater fish- es of Canada. Fisheries Research Board of Canada Bulletin Number 184. Ottawa, Ontario. 996 pages. Received 19 December 2001 Accepted 18 January 2003 Over-wintering and Reproduction by the Big Brown Bat, Eptesicus fuscus, in New Brunswick: DONALD F. McALpINe!, FRANCES MULDOON”, GRAHAM J. FoRBES?, ALEXANDER I. WANDELER2, ScoTT MAKEPEACE’, HuGH G. BRopERS?, and JAMES P. GoLTz> ‘Natural Science Department, New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada; e-mail: dmcalpin@nb.aibn.com *Canadian Food Inspection Agency, Animal Diseases Research Institute, 3851 Fallowfield Road, P.O. Box 11300, Station H, Nepean, Ontario K2H 8P9 Canada ’New Brunswick Cooperative Fish and Wildlife Research Unit, University of New Brunswick, Fredericton, New Brunswick E3B 6C2 Canada *Fish and Wildlife Branch, New Brunswick Department of Natural Resources and Energy, P.O. Box 6000, Fredericton, New Brunswick E3B 5H1 Canada Provincial Veterinary Laboratory, P.O. Box 6000, Fredericton, New Brunswick E3B 5H1 Canada McAlpine, Donald F., Frances Muldoon, Graham J. Forbes, Alexander I. Wandeler, Scott Makepeace, Hugh G. Broders, and James P. Goltz. 2002. Over-wintering and reproduction by the Big Brown Bat, Eptesicus fuscus, in New Brunswick. Canadian Field-Naturalist 116(4) : 645-647. The first evidence of over-wintering and reproduction by the Big Brown Bat, Eptesicus fuscus, in New Brunswick is docu- mented based on 24 new specimens and other reports from southern New Brunswick. Previously, this species was known in New Brunswick from three specimens and several unverifiable sightings. New distributional records, combined with echolocation surveys, suggest that E. fuscus is an insignificant component of forested ecosystems along the Fundy coast of central New Brunswick. In New Brunswick, the Big Brown Bat appears to be closely associated with human-occupied buildings. New Brunswick data support a previous suggestion that such behaviour may have promoted a northward range expansion by E. fuscus, and perhaps an increase in populations in more northerly regions. Key Words: Big Brown Bat, Eptesicus fuscus, behaviour, distribution, hibernation, reproduction, New Brunswick The Big Brown Bat, Eptesicus fuscus, reaches the northern limit of its eastern North American range in southern New Brunswick (van Zyll de Jong 1985). Although Morris (1948) stated that the Big Brown Bat occurred in New Brunswick, the presence of E. fuscus in the province was not confirmed until 4 September 1959 when a specimen was collected at St. Andrews, Charlotte County (Gorham and Johnston 1962). Squires’ (1968) synopsis for the species in the province included only this single specimen and an indirect reference to an observation by S. W. Gorham. Subsequently, two additional specimens have been collected at Saint John, St. John County, 3 September 1976 and near Mactaquac, York County, on 4 September 1982. There is also a report of a Big Brown Bat echoloca- tion sequence from Fundy National Park. (Tremblay 1989*, 1992) and several unsubstantiated sight * see Documents Cited section records from various southern New Brunswick loca- tions dating from 1930 to 1995 (Appendix 1; Squires 1967; Christie 1986). Sight records are based on bats observed roosting in a barn, in flight, or removed from mist nets during bird banding operations. Until now, there has been no indication that the Big Brown Bat reproduced in New Brunswick. Also, in spite of extensive searches of New Brunswick bat hibernacula in caves and abandoned mines there have been no observations (McAlpine 1983) of this bat species over-wintering in the province. Here we compile 39 reports, both published and unpublished, for the Big Brown Bat in New Brunswick. These reports include 24 new specimens, plus two addition- al observations verified with archived photos and measurements. Most of the new data presented here has been collected incidental to ongoing monitoring programs for rabies. There are five cases of the Big Brown Bat over- wintering in New Brunswick. Four of these bats, and 646 possibly the fifth, were all discovered moving about inside heated buildings between December and 23 March 1996-2001; NBM files, NBM 5796, 5821, 5936). Among the 24 new specimen records is a non- volant, near hairless, young collected from a building in Fredericton, 22 June 1999 (NBM 5792). A home in Central Hampstead, Queens County, also appeared to support a maternity colony under the metal flashing covering the roof of a 19" century farmhouse. One of the bats (NBM 5797) taken at this site, although well- furred, has a forearm length of 36.1 mm well below the range for adults of the species cited by van Zyll de Jong (1985). Until recently, E. fuscus was known primarily as a cave or mine hibernator. However, Whitaker and Gummer (2000) suggest that prior to European colonization of North America, the Big Brown Bat hibernated mainly in hollow trees, and that this has pre-adapted the species to exploit buildings. New evidence shows that the hibernacula of choice for this species in Indiana are the attics of heatéd build- ings (Whitaker and Gummer 1992; Whitaker and Gummer 2000). Whitaker and Gummer (2000) pro- pose that historically the Big Brown Bat was limited to a more southerly range due to a dependence on hollow trees for hibernation, but recent use of heated buildings may have allowed the species to extend its range northward and to increase populations in more northerly areas. However, Whittaker and Gummer (2000) found that most buildings serving as hiber- nacula harboured very few bats (x = 2.7 bats for buildings which did not serve as summer maternity roosts). Brigham (1987) has reported that E. fuscus discovered active in buildings during winter in Ottawa weighed less and had significantly shorter forearm lengths than inactive bats hibernating in caves. The short mean forearm length of active bats studied by Brigham (1987) indicates that many were juveniles. He hypothesized that E. fuscus may become active in winter when energy reserves are critically low. While this suggests that active bats may not be those which will survive the winter in these buildings, two of our records came from the same building five years apart, suggesting long-term use of the site and successful over-wintering by some individuals. Records presented here provide the first evidence of E. fuscus as a reproducing and over-wintering species in New Brunswick. While uncommon, the Big Brown Bat appears to be more abundant in New Brunswick than documented previously. With few Big Brown Bats in buildings, it is possible that the species has been overlooked in the past in New Brunswick. However, there have been an increasing number of bats submissions from New Brunswick for rabies testing since 1987 (6 in 1987, a high of 48 in 2001), the source for most of the recent Big Brown Bat data for the province presented here. THE CANADIAN FIELD-NATURALIST Vol. 116 - Only about 30 echolocation sequences from nearly 160 000 collected in and around Fundy National Park show characteristics that suggest these sequences are either Big Brown or Silver Haired Bat (Lasionycteris noctivagans) (H. Broders, University of New Brunswick, unpublished). Likewise, bat sur- veys in central forested and southwestern coastal Nova Scotia show the Big Brown Bat to be virtually absent in these areas (Broders et al. in review*). Also, Zimmerman and Glanz (2000) captured only four E. fuscus in 840 mist net hours along the forest- ed central Maine coast. While it appears that E. fus- cus is an insignificant component of forested ecosys- tems along the central Fundy coast of New Brunswick, targeted surveys for this species are lack- ing. In particular, there is a need to carry out echolo- cation surveys in tolerant hardwood stands in south- western New Brunswick, where large trees (> 50 cm DBH) that might provide rooting sites are present, or at likely feeding areas nearby. In New Brunswick the Big Brown Bat reaches a northern range limit, where it appears to be closely associated with human-occu- pied buildings. This supports the suggestion by Whittaker and Gummer (2000) that such behaviour may have promoted a northward range expansion by E. fuscus, and perhaps an increase in populations in more northerly regions. Acknowledgments Mark Phinney and Dwayne Sabine, University of New Brunswick Forestry Department, generously provided details on one of the over-wintering records from a building on the University of New Brunswick, Fredericton campus. Comments by Fred Scott and an anonymous referee improved the manuscript. Documents Cited (marked * in text) Tremblay, E. 1989. A brief survey of the chiropterian fauna of Fundy National Park. Unpublished report, Parks Canada, Natural Resources Conservation, Fundy National Park. 19 pages plus appendices. Broders, H. G., G. M. Quinn, and G. J. Forbes. (in review). Species status, and the spatial and temporal pat- terns of activity of bats in southwest Nova Scotia, Canada. Northeastern Naturalist. Literature Cited Brigham, R. M. 1987. The significance of winter activity by the Big Brown Bat (Eptesicus fuscus): the influence of energy reserves. Canadian Journal of Zoology 65: 1240-1242. Christie, D.S. 1986. [Big Brown Bat on Grand Manan]. Nature News — Autumn 1985. New Brunswick Naturalist 15: 35. Gorham, S. W., and D. H. Johnston 1962. Notes on New Brunswick bats. Canadian Field-Naturalist 76: 228. McAlpine, D. F. 1983. Status and conservation of solu- tion caves in New Brunswick. New Brunswick Museum Publications in Natural Science Number 1. 28 pages. Morris, R. F. 1948. The land mammals of New Brunswick. Journal of Mammalogy 29: 165-176. 2002 Squires, W. A. 1967. [Big Brown Bat at Westfield]. Nature News (New Brunswick Museum, Saint John) 18: 3. Squires, W. A. 1968. The Mammals of New Brunswick. Monographic Series (5). New Brunswick Museum, Saint John, 57 pages. Tremblay, E. 1992. Bats of Kouchibouguac and Fundy National Parks, New Brunswick, Canada. Pages. 291-294 in Science and the Management of Protected Areas. Edited by J. Willison, S. Bondrup-Nielson, C. Drysdale, T. B. Herman, N. W. P. Munro, and T.L. Pollock. Elsevier, Amsterdam. van Zyll de Jong, C. G. 1985. Handbook of Canadian Mammals (2): Bats. National Museum of Natural Sciences, Ottawa. 212 pages. NOTES 647 Whitaker, J. O., Jr., and S. L. Gummer. 1992. Hibernation of the Big Brown Bat, Eptesicus fuscus, in buildings. Journal of Mammalogy 73: 312-316. Whitaker, J. O., Jr., and S. L. Gummer. 2000. Population structure and dynamics of Big Brown Bats (Eptesicus fuscus) hibernating in buildings in Indiana. American Midland Naturalist 143: 389-396. Zimmerman, G. S. and W. E. Glanz. 2000. Habitat use by bats in eastern Maine. Journal of Wildlife Management 64: 1032-1040. Received 27 February 2001 Accepted 28 January 2003 Appendix 1. Date, location, and source for records for the Big Brown Bat, Eptesicus fuscus, in New Brunswick. *Bats verifiable by a specimen, or a photograph accompanied by measurements, deposited in the New Brunswick Museum (NBM) or Canadian Museum of Nature (CMN) collections. ' Bats submitted to Animal Disease Research Institute, Ottawa for rabies testing. Big Brown Bats submitted for testing prior to 1996 cannot be verified with specimens; however, at least four of the five animals were identified by a technician experienced in identifying Canadian bats. Minimally, each of these records is documented in files housed at the NBM. July 1930, Browns Flat, Kings County, NBM files. *4 September 1959, nr. St. Andrews, Charlotte County, CMN 27731 October 1967, Westfield area, Kings County, Squires (1967) *3 September 1976, Saint John, St. John County, NBM 1347 *4 September 1982, Mactaquac, York County, NBM 1887 1985!, Moncton , Westmorland County, NBM files 27 October 1985, Grand Manan, Charlotte County, Christie (1986) 23 June 1987!, Moncton, Westmorland County, NBM files 11 August 1987! , Fredericton, York County, NBM files 21 August 1988, Fundy National Park, Albert County, Tremblay (1989) 20 July 1988', Sackville, Westmorland County, NBM files *? February 1990, Fredericton, York County, NBM files 12 July 1991', Fredericton, York County, NBM files 25 September 1995, Kent Island, Charlotte County, NBM files #15 July 1996!, Fredericton, York County, NBM 5933 *December 1996, Fredericton, York, County, NBM files *24 July 1997, Central Hampstead, Queens County, NBM 5793 *24 July 1997', Central Hampstead, Queens County, NBM 5794 *24 July 1997', Central Hampstead, Queens County, NBM 5795 *24 July 1997', Central Hampstead, Queens County, NBM 5797 *15 January 1998', St. Andrews, Charlotte County, NBM 5796 *6 August 1998', Saint John, Saint John County, NBM 5825 *22 June 1999!, Fredericton, York County, NBM 5792 * 2 July1999', Central Hampstead, Queens County, NBM 5823 * 2 July1999', Central Hampstead, Queens County, NBM 5935 *14 July 1999!, Fredericton, York County, NBM 5791 *15 February 2000', Fredericton, York County, NBM 5821 *18 July 2000', Fredericton, York County, NBM 5820 * 30 August 2000', Lincoln, York County, NBM 5819 *23 March 2001', Fredericton, York County, NBM 5936 *§ June 2001', Harvey, York County, NBM 5937 *20 June 2001! , Islandview, York County, NBM 5938 *25 June 2001', St. Stephen, Charlotte County, NBM 5939 *29 June 2001', Central Hampstead, Queens County, NBM 5934 *20 August 2001', St. Stephen, Charlotte County, NBM 5940 *4 September 2001', Moncton, Westmorland County, NBM 5951 *§ August 2002', Sussex, Kings County, NBM 5952 *14 August 2002', Fredericton, York County, NBM 5953 *21 August 2002', Black's Harbour, Charlotte County, NBM 5954. —— hhh eee, ne 648 THE CANADIAN FIELD-NATURALIST Vol. 116 - Behavioral Modification of Gray Wolves, Canis lupus, Suffering from Sarcoptic Mange: Importance of Sequential Monitoring DouGLas P. SHELLEY! and THOMAS M. GEHRING2 \Department of Agricultural Sciences, University of Wisconsin-River Falls, River Falls, Wisconsin 54022 USA 2Department of Biology, Central Michigan University, Mt. Pleasant, Michigan 48859 USA Shelley, Douglas P., and Thomas M. Gehring. 2002. Behavioral modification of Gray Wolves, Canis lupus, suffering from sarcoptic mange: Importance of sequential monitoring. Canadian Field-Naturalist 116(4): 648-650. We documented the behavioral patterns of two disease-stricken Gray Wolves (Canis lupus) in northwestern Wisconsin and east-central Minnesota by means of radio telemetry. Our observations suggest that the modified behaviors of these wolves were integrally linked to the increased loss of thermal energy due to severe hair loss and sarcoptic mange mite and biting louse infestations. We believe that sequential monitoring using radio telemetry allowed the identification of these behav- ioral patterns. Wildlife managers might use frequent monitoring to identify disease-stricken animals and implement proac- tive measures to counter some wildlife diseases. Key Words: Canis lupus, Gray Wolf, disease, behavior, mortality, radio telemetry, sarcoptic mange, Wisconsin, Minnesota. Radio telemetry is a useful technique for gathering data on wide-ranging and cryptic wildlife species. Recently, several researchers have debated the ques- tion of independence in radio locations (Swihart and Slade 1985a, 1985b, 1997). Swihart and Slade (1997) reiterated the importance of independent radio locations (e.g., for some habitat selection analyses), but also pointed to the importance of sequential observations for behavioral studies. Herein, we report on the modified behavior of Gray Wolves (Canis lupus) suffering from sarcoptic mange as determined by means of sequential moni- toring. We suggest that frequent monitoring may assist managers in the early detection of disease in recovering wolf populations. Methods Wolves were captured in east-central Minnesota and northwestern Wisconsin during May-August in modified #14 Newhouse steel traps (Kuehn et al. 1986). Captured wolves were immobilized by means of an intra-muscular injection of ketamine hydrochloride and xylazine hydrochloride. Once immobilized, wolves were fitted with a radio-collar and ear-tagged with a uniquely numbered plastic tag (Kohn et al. 2000). During October 1992-January 1993, we monitored the movements of individuals from the Truck Trail pack (TTP) using a vehicle- mounted radio telemetry unit (Gehring 1995; Shelley and Anderson 1995). The TTP consisted of four wolves, three of which were radio-tagged, including: 131M (adult male, captured 21 August 1991, capture weight = 44.9 kg), 203F (adult female, captured 29 July 1992, capture weight = 30.8 kg), and 133F (yearling female, captured 22 August 1991). Wolf 133F dispersed from the TTP in December 1992. The body weight of W203F was comparable to the mean body weight for adult female wolves in our *see Documents Cited study area (x= 30.2 kg, n= 4; Gehring 1995), where- as the body weight of W131M was greater than the mean body weight for adult male wolves in our study area (x= 37.1 kg, n = 7; Gehring 1995). Only wolves 203F and 133F radio collars were equipped with mortality sensors (Garshelis et al. 1982). Wolves 203F and 131M were relocated by triangula- tion (Heezen and Tester 1967) 1-2 times daily and hourly during sequential tracking sessions. Radio locations obtained > 4 h apart were considered inde- pendent (Swihart and Slade 1985a, 1985b, 1997). Sequential tracking sessions were conducted for 6-8 hours, approximately 4-6 times/month. Additionally, we recorded the activity of wolves by noting changes in pulse rates (Garshelis et al. 1982). Results and Discussion During the study period, we collected 107 and 102 independent locations for 203F and 131M, respec- tively. The time between locations ranged from 4-48 h (203F) and 4-120 h (131M). We relocated these two wolves together during 24% of the October loca- tions, 8% of the November locations, 5% of the December locations, and none of the January loca- tions. Conversely, we found that wolves maintained high pack cohesion in three neighboring packs that we monitored concurrently (Gehring 1995). We found that both wolves concentrated their move- ments to four areas within the TTP territory (33% of 203F and 37% of 131M locations), but these individ- uals largely used these areas independently of one another. Two of these sites contained carrion; we found a Beaver (Castor canadensis) and Mink (Mustela vision) carcass pile left by trappers at one site, and two to three carcasses of White-tailed Deer (Odocoileus virginianus) killed by wolves in December at the other site. We recorded numerous visits and wolf bedding areas at the latter site. The third site contained no obvious food source, and the fourth was considered a non-food area until January 2002 when the remains of a freshly-killed White-tailed Deer were found. We believe that 203F made this kill with another wolf based on wolf tracks present and our telemetry record. From 23 December 1992 to 13 January 1993, 203F exclusively traveled a network of improved dirt roads, plowed and unplowed. A total of 26 indepen- dent locations were obtained during this period. Of these, 5 (19%) were recorded in mortality mode (i.e., the animal had not moved in 5-6 h). After 203F was initially located, she would remain in mortality mode for an additional 4-27 h (determined by sequential locations). On 13 January 1993, 203F was located in a local resident’s garage. She was immobilized and transported to a local wildlife health clinic for ectoparasite treatment on 14 January, but died later that night. Necropsy results indicated that the primary cause of death was a subcutaneous fistula leading to pyothorax (Thomas 1993a*). An abscess in the mus- cles of the right thoracic region was centered around an embedded Porcupine (Erethizon dorsatum) quill and was estimated to be several weeks old. The Porcupine quill may have become embedded in 203F if she was scavenging or rolling on a Porcupine car- cass (Mech 1973), or the remote possibility of her preying on the Porcupine (not found in the literature). Wolf 203F was suffering from severe infestations of sarcoptic mites (Sarcoptes scabiei var. canis) and bit- ing louse (Trichodectes canis) and approximately 35% hair loss (Thomas 1993a*), indicating moderate- ly severe alopecia (Bornstein et al. 2001). Wolf 203F’s carcass weight, 27.4 kg, was 11% less than her capture weight. From 1-26 January 1993, the 24 locations obtained on 131M indicated that he restricted his movements to an approximately 5 ha area. Unfortunately his collar was not equipped with a mortality sensor. However, 15 of 131M’s January locations were recorded in inactive mode (Kenward 1987), but he was believed to move between loca- tions due to noticeably different UTM locations. Radio locations suggested that 131M did not move from 22-26 January 1993 (no discernable UTM dif- ference). Wolf 131M was found dead on 27 January 1993, buried under 20-25 cm of snow. Wolf 131M’s time of death was estimated to be between 20-26 January 1993 based on snow fall data (snow fall recorded on 12-14 January and 22-26 January 1993) and telemetry locations. Necropsy diagnostic results indicated cold exposure as the primary cause of death (Thomas 1993b*). W131M had severe infesta- tions of sarcoptic mange mites and biting lice. Approximately 50-75% of 131M’s body hair was missing (Thomas 1993b*), indicating severe alope- cia (Bornstein et al. 2001). Wolf 131M’s carcass weight was 43.0 kg, a 4% decrease in weight from his capture date. Todd et al. (1980) documented that mangy wolves in Alberta had body weights 4-10% lower than non- NOTES 649 mangy wolves. Wolves with advanced cases of mange were 22% below the average weight for adults (Todd et al. 1980). In terms of body condition (based on loss of body weight), W203F appeared to be in the poorest body condition resulting from a moderately severe case of mange, whereas W131M appeared to have slightly better body condition based on a 4% loss of weight. We observed behavioral patterns in wolves 203F and 131M which appear to differ from wolf behavior stereotyped as normal behavior (sensu Mech 1970). Both collared wolves appeared to increase scavenging behavior during October-January. For wolves, behav- ioral shifts to increased scavenging are generally asso- ciated with low prey availability or illness (Mech 1973; Todd et al. 1980). However, deer and Beaver abundance was high in the TTP (Gehring 1995; Shelley and Anderson 1995). Trainer and Hale (1969) observed mangy Coyotes (Canis latraus) that demon- strated abnormal behavior including listlessness and loss of fear of man. Similarly, Todd et al. (1980) sug- gested that mangy coyotes tended to be more depen- dent on carrion compared to non-mangy coyotes. Thus, the apparent increased use of carrion by the two wolves in our study suggests that mange was the causal factor leading to these increased scavenging. Increased dependence on carrion, concentrated movements, extended periods of inactivity (by both wolves), immobility defined by the mortality sensor, and exclusive travel on roads suggests that these wolves were in poor physical condition. We suggest that this weakened physical state and modified behavior was integrally linked to the increased loss of thermal energy due to severe hair loss and sarcop- tic mange mite and biting louse infestations. We believe that sequential observations allowed us to identify these behavioral modifications. Furthermore, we suggest that managers might use sequential radio locations and frequent monitoring to define dramatic shifts in behavioral patterns, particu- larly in recovering populations (e.g., Wisconsin Department of Natural Resources 1999*). Such shifts may be used as an indicator of disease-stricken animals and may allow proactive treatment of wildlife diseases. Acknowledgments Funding and logistical support was provided by the Wisconsin Department of Natural Resources (Federal Aid in Wildlife Restoration Act, Pittman- Robertson Project W-141-R); Wisconsin Department of Transportation; and University of Wisconsin- Stevens Point. We also thank the Minnesota Department of Natural Resources, Douglas County Forestry Department, and U. S. Fish and Wildlife Service for additional logistical support of this study, E. M. Anderson and B. E. Kohn provided valuable comments on various drafts of this manuscript. 650 Documents Cited (marked * in the text) Kohn, B., J. Frair, D. Unger, T. Gehring, D. Shelley, E. Anderson, and P. Keenlance. 2000. Impacts of the U.S. Highway 53 expansion project on wolves in north- western Wisconsin: final report. Wisconsin Department of Natural Resources. 49 pages. Shelley, D. P., and E. M. Anderson. 1995. Effects of vehicular traffic on and ecology of timber wolves in northwestern Wisconsin and east-central Minnesota. Final Report. University of Wisconsin-Stevens Point, Stevens Point. Thomas, N. J. 1993a. National Wildlife Health Research Center Necropsy Report. U. S. Fish and Wildlife Service National Health Research Center. Madison, Wisconsin. #11249. Thomas, N. J. 1993b. National Wildlife Health Research Center Necropsy Report. U. S. Fish and Wildlife Service National Health Research Center. Madison, Wisconsin. #11291. 5 Wisconsin Department of Natural Resources. 1999 Wisconsin wolf management plan. Madison, Wisconsin. PUBL-ER-099 99. 74 pages. Literature Cited Bornsten, S., T. Moérner, and W. M. Samuel. 2001. Sarcoptes scabiei and sarcoptic mange. Pages 107-119 in Parasitic diseases of wild mammals. Edited by W. M. Samuel, M. J. Pybus, and A. A. Kocan Iowa State University Press, Ames, Iowa. Garshelis, D. L., H. B. Quigley, C. R. Villarrubia, and M. R. Pelton. 1982. Assessment of telemetric motion sensors for studies of activity. Canadian Journal of Zoology 60: 1800-1805. Gehring, T. M. 1995. Winter wolf movements in north- western Wisconsin and east-central Minnesota: a quanti- tative approach. M. S. thesis, University of Wisconsin- Stevens Point, Stevens Point, Wisconsin, USA. Heezen, K. L., and J. R. Tester. 1967. Evaluation of radio- tracking by triangulation with special reference to deer movements. Journal of Wildlife Management 31: 124-141 THE CANADIAN FIELD-NATURALIST —L— Se LAs Vol. 116 “ Kenward, R. 1987. Wildlife Radio Tagging: equipment, field techniques, and data analysis. Academic Press, New York, New York, USA. Kuehn, D. W., T. K. Fuller, L. D. Mech, W. J. Paul, S. H. Fritts, and W. E. Berg. 1986. Trap-related injuries to gray wolves in Minnesota. Journal of Wildlife Management 50: 90-91. Mech, L. D. 1973. The wolf: the ecology and behavior of an endangered species. University of MN Press, Minneapolis, Minnesota, USA. Mech, L. D. 1977. Productivity, mortality and population trends of wolves in northwestern Minnesota. Journal of Mammalogy 58: 559-574. Schwartz, C. C., R. Stephenson, and N. Wilson. 1983 Trichodectes canis on the Gray Wolf and Coyote on Kenai Peninsula, Alaska. Journal of Wildlife Diseases 19: 372-373. Swihart, R. K., and N. A. Slade. 1985a. Influence of sampling interval on estimates of home range size. Journal of Wildlife Management 49: 1019-1025. Swihart, R. K., and N. A. Slade. 1985b. Testing for inde- pendence of observations in animal movements. Ecology 66: 1176-1184. Swihart, R. K. and N. A. Slade. 1997. On testing for independence of animal movements. Journal of Agricultural, Biological, and Environmental Statistics 2: 48-63. Todd, A. W., J. R. Gunson, and W. M. Samuel. 1980 Sarcoptic mange, an important disease of coyotes and wolves of Alberta. Pages 706-729. in Proceedings of the first worldwide furbearer conference. Edited by J. Chapman and D. Pursley. Frostberg College, Frostberg, Maryland, USA. Trainer, D. O., and J. B. Hales. 1969. Sarcoptic mange in red foxes and coyotes of Wisconsin. Bulletin of the Wildlife Disease Association 5: 387-391. Received 29 June 2001 Accepted 11 October 2002 Book Reviews ZOOLOGY The Birds of British Columbia Volume 4: Passerines [:] Wood-Warblers through Old World Sparrows By R. Wayne Campbell, Neil K. Dawe, Ian McTaggart- Cowan, John M. Cooper, Gary W. Kaiser, Andrew C. Stewart, and Michael C. E. McNall. 2001. UBC Press, Vancouver, British Columbia. 741 pp., illus. $125.00. - This hefty tome completes a series inaugurated in 1990 (see reviews of the first two volumes in Canadian Field-Naturalist 107: 547-548, 1993 and of volume 3 in Canadian Field-Naturalist 115: 187- 188, 2001). The bulk of the book (pp. 9-617) consists of accounts of species not covered in the first three volumes. These cover 101 species in seven passerine families (Parulidae: wood-warblers, Thraupidae: tanagers, Emberizidae: New World sparrows and allies, Cardinalidae: some grosbeaks, some buntings and Dickcissel, Icteridae: blackbirds, orioles and allies, Fringillidae: Cardueline finches and allies, and Passeridae: Old Word Sparrows). A second major section (pages 619-629) consists of mini accounts of 28 species that have been docu- mented as occurring within British Columbia or as breeding within the province since the publication of the first two volumes (non-passerines), not the first three as stated on page 620. A final major section consists of a two-chapter synopsis (pages 631-695). Additional introductory and closing sections include a detailed, four-page acknowledgement of significant contributions as well as a six-page list of 3629 (!) contributors of records, a list of references cited, a brief index, and brief biographical details of the authors and of four additional “contributing” authors, who wrote sections of volume | (Dennis A. Demarchi) or of this volume (Richard R. Howie, Chris Siddle, and Linda M. Van Damme). Four data appendices were omitted from the book for space reasons, but are available on the World Wide Web for interested readers who have access to the inter- net. A brief tribute by his co-authors to Ian McTaggart-Cowan’s contributions to British Columbia ornithology over his first 90 years con- cludes the book. As in the first three volumes, species of “regular” occurrence receive more coverage than in many other regional avifaunal works of recent decades. These accounts encompass three to 17 pages each, with text on range (generally in the world), status in British Columbia, nonbreeding and breeding status, distribution and chronology in the province, nesting details if the species breeds in British Columbia (with details on nests, eggs, young, degree of Brown-headed Cowbird parasitism [if any]), “remarks” (comments on taxonomy, range expan- sions/contractions, comments on questionable records, remarks on range and distribution of distinct races within the province, and numerous other bio- logical tid-bits), and a selection of “noteworthy” records. The text is illustrated with colour pho- tographs of the species, its habitat(s) in British Columbia, and sometimes nests, eggs, and/or young and with maps and graphs of distribution and chronology in various regions of this vast province with its complex range of habitats and climate. Additional graphs, maps, and tables in some species illustrate population trends, chronology of nesting in different regions, banding recoveries, a tabulation of known hosts of Brown-headed Cowbirds in British Columbia and other biological details. Relevant liter- ature is cited throughout the text. As in the previous volumes, species considered “casual, accidental, extirpated and extinct” in the province receive shorter accounts (about one-third to one page each). These 22 accounts consist of the world range of the species, its status in British Columbia, a list of occurrences accepted by the authors, and “remarks”. Documentary photographs accompany ten of these accounts. The authors’ desire to be as current as possible is exemplified by the addition in postscripts of occurrences of three of these species after the species accounts were written. The section of addenda to non-passerine birds consists of brief accounts of six species (Clark's Grebe, Northern Fulmar, Upland and Baird’s sand- pipers, Franklin’s Gull, and Black-legged Kittiwake) demonstrated to breed in British Columbia since publication of the 1990 volumes, as well as accounts of one species added to the province’s avifauna by virtue of a taxonomic split (Pacific Golden-Plover), four species elevated from “hypothetical” to docu- mented status (Black Vulture, Falcated Duck, Spectacled Eider, and Yellow Rail), and 17 species completely new to the provincial list. The concluding “synopsis” consists of two chap- ters. The first, “avian biodiversity, ecological distri- bution, and patterns of change” is essentially a condensed analysis of ornithogeography in British Columbia, examining patterns of distribution (breed- ing, wintering, and migrant) in relation to geographic features, human activity, and other factors. These 651 652 emphasize changes in both distribution and popula- tions in relation to various environmental changes. Such concepts as biodiversity, species richness, “hot spots” in which large numbers of birds concentrate, endemism, population trends, and species of varying degrees of concern are discussed with the aid of a rich array of tables and graphs. The final chapter is a realistically pessimistic look at the future of bird populations in British Columbia and the world gen- erally, summarizing the role of human activity of all sorts (depletion of resources, altering of ecosystems, etc.) in putting ever-increasing stresses upon the environments on which birds and other life forms depend as human populations grow and the impacts imposed by individual humans escalate. Local and global solutions required to counteract these trends are also outlined. The vast geographic expanse of British Columbia and its rich variety of habitats at varying elevations pose challenges to writing generalized accounts of birds that occur there regularly. The authors have handled this diversity well by noting status within various portions of the province, and charting such features as migration and egg-laying dates by region in text and/or in charts. Although knowledgeable readers of any specific account will quibble with omissions of specific details and/or references, the text is both more thorough than is usual in provincial or state bird books and well referenced. Emphasis is on British Columbia data when available, but specif- ic details from elsewhere are cited frequently. On the whole, the accounts of species that have strayed into British Columbia are also thorough and written well. When available, documentary pho- tographs are included. The text places the British Columbia occurrence(s) in context with population trends and patterns of vagrancy elsewhere. The “remarks” sections of these species often include notes on additional records of the species that the authors have rejected because of lack of sufficient documentation or suspicions that the bird observed had escaped from captivity. Inclusion of such records will help observers assess future records of these species, some of which may seem more likely as patterns of vagrancy become apparent. Unfortunately, however, disagreement on the validi- ty of records accepted by the authors is not men- tioned. For example, a Xantus’s Hummingbird that spent nearly a year at Gibsons is included without any mention that the record was rejected by the THE CANADIAN FIELD-NATURALIST Vol. 116 British Columbia Field Ornithologists Bird Records Committee on the grounds that it may represent a ship-transported bird. While many observers (including several of the committee members) agree with the authors in accepting this record, it would be useful to readers to know that the record is controversial. Similarly, no mention is made that one of the three records of White-tailed Kite accepted by the authors was rejected by the B.C. records committee, but accepted by the equivalent Vancouver committee. The chapter on “avian biodiversity, ecological distribution, and patterns of change” is a thorough summary of distributional patterns and ongoing pop- ulation and distributional changes within British Columbia of all the species covered in all four volumes. The text is enhanced by numerous tables, maps, and charts. Canadian Field-Naturalist readers will already be familiar with the many threats to bird populations (and ecosystems generally) outlined in the final chapter. As in previous volumes, the list of references cited is extensive. Unfortunately, about 30-40 of the refer- ences cited in the text appear not to be in the list of references. Some of these probably result from mis- matches between the dates indicated in the text and the dates of the corresponding listed references. The number of missing or mismatched references seems high for a book with so many authors. The final proof-reading appears not to have been performed by all of the authors. Perhaps the most significant omis- sion is a reference by Stone to a male Sage Sparrow collected in Vancouver in 1899 and overlooked in previous works on British Columbia’s avifauna. As in volume 3, the 1966 edition of The birds of Alberta by W. R. Salt and A. L. Wilk is listed as by W. R. and J. R. Salt. Most of the errors in this volume consist of minor grammatical flaws (especially split infinitives), spelling of the names of some observers, and other minor glitches, rather than substantial mistakes. Like the three previous volumes, this final volume is a thorough compilation and analysis of what is known about British Columbia’s avifauna to date and an excellent guide to many of the knowledge gaps that still exist. MARTIN K. MCNICHOLL 4735 Canada Way, Burnaby, British Columbia V5G 1L3 Canada 2002 BOOK REVIEWS 653 The Royal Ontario Museum Field Guide to Amphibians and Reptiles of Ontario By Ross MacCulloch. 2002. Royal Ontario Museum and McClelland and Stewart Ltd., Toronto, Ontario. 168 pp., illus. $24.99 Ontario, with more than half of the over 90 species (native and introduced) of amphibians and reptiles recorded in Canada, was the first province to have its own guides to its amphibians (1937, Royal Ontario Museum Handbook 3) and reptiles (1939, Royal Ontario Museum Handbook 4). The author of these, E. B. S. Logier, artist and herpetologist at the ROM from 1915 to 1961, followed with books on amphibians of eastern Canada (1952, Clarke, Irwin & Company Limited) and snakes of Ontario (1958, University of Toronto Press). Since, there have been popular booklets on Ontario snakes and turtles in 1967 by Barbara Froom published by the Ontario Department of Lands and Forests (now Ministry of Natural Resources) and in 1989 a book, Familiar Amphibians and Reptiles of Ontario by Bob Johnson of the Metropolitan Toronto Zoo published by Natural Heritage/National Heritage Inc., Toronto. Detailed spot distribution maps for the province have been underway as the Ontario Herpetofaunal Summary (1986, 1988, 1989) edited by Michael J. Oldham and others, and now updated on a web site sponsored by the Natural Heritage Information Centre of the Ontario Ministry of Natural Resources www.mnr.gov.on.ca/NMR/nhic/herps/ohs.html. Now comes a notching up in the standards of pre- sentation by several orders of magnitude. The ROM Field Guide to Amphibians and Reptiles of Ontario is a blaze of colour photographs offset with brief and usefully structured information, truly a field-natural- ist's book. What it lacks in detailed character or eco- logical discussions prevalent in earlier Ontario guides, it offsets by providing ease of identification through colour photographs and pruned accounts most useful for field use. The experience of the author is evident throughout. MacCulloch, now an assistant curator, has been active in herpetology at the Royal Ontario Museum since 1982 with research projects both in Ontario and abroad, and, previous to that, has conducted field studies in Quebec with Roger Bider of Macdonald College, McGill University, and obtained a M.Sc. on the Western Painted Turtle in Saskatchewan with Diane Secoy, University of Regina. The organization and layout of the book are appealing. A brief Introduction sets amphibians and reptiles in context in natural systems. Characteristics covered are general features of reproduction, metamorphosis, skin, diet, and metabolism for amphibians and reproduction, eggs, skin, diet, and metabolism for reptiles. Treated next are The Challenge of Climate (winter, hibernacula, spring, summer and autumn); Observing Amphibians (oddly omitting frogs); Identification (appearance, habitat, season, adding to our understanding — with reference to data bases and surveys which need volunteers, and naturalists groups which conduct nature walks and field trips); Conservation (legal issues, amphibians and reptiles as pets — pointedly stressing that the reg- ulations that exist on keeping native species as well as the difficulties with maintaining exotics argue against captives); and Amphibian populations in decline? — stressing their general resilience despite cases of localized extirpations of populations or species). Using This Book discusses the format of sections to follow on Comparative Photographs and of Species Accounts (and includes national (COSEWIC) and Ontario (OMR) species ranking definitions), and Distribution Maps. Further Information on Ontario Amphibians and Reptiles contains a list of selected books and other print sources and a list of internet sources which cover provincial and national parks in Ontario, other agen- cies and organizations, and Ontario natural history societies (even if you are not the least interested in amphibians and reptiles this is a handy reference for any naturalist). The book ends with a glossary of terms, a “Checklist of Ontario amphibians and rep- tiles”, index, photo credits, and acknowledgments. The bulk of the book lies in the user-friendly com- parative photographs section (pages 25-32) and the species accounts (pages 25-32). The comparative photographs, effectively replacing written keys used in many other publications, are grouped by salaman- ders, frogs, turtles, and snakes (the one lizard, the only representative of its group, is omitted here) with one photograph and the Ontario distribution map in miniature for each species for quick contrast. The species accounts proper are equally adroitly designed and cover the 47 species (10 salamanders, 13 frogs, 8 turtles, 1 lizard, and 15 snakes) regarded as sull occurring and native. For each species a page of three colour photographs of variants in colour, pat- tern, or life history stage is opposite a page giving introductory comment, appearance, habitat and behaviour, reproduction, and status. There is also a map of Ontario, and one of North American, distri- bution for each. The ranges are depicted as solid patches and the reader is directed to the Ontario Herpetofaunal Survey web posting for more detailed distribution maps, a reasonable approach assuming most field users will want approximate rather than detailed distributions (which would not work well in the pocket-sized format of the book). In addition to the full accounts, page 96 briefly gives a paragraph each to three salamanders and page 162 to two tur- tles and one snake regarded as extirpated (Tiger Salamander, Spring Salamander, and Timber Rattlesnake) or introduced (Red Salamander, Box Turtle, Red-eared Slider) to bring the total recorded 654 for Ontario to 53). Amphibians and reptiles as groups are each given, before the accounts of their included species, two pages outlining general char- acteristics. Each family within these is prefaced by a photographic portrait of a member of that group opposite a paragraph of text on group characteristics. An added convenience is a colour strip on the edge of each page of the comparative photographs and species accounts with topic captions along them, so the reader can flip the edges and rapidly find the account desired. There are a few niggling details amiss to irritate the experienced herpetologist, inevitable in a book where the text is so concentrated and photographs are from varied contributors. Among omissions are the lack of specific mention in the Blue-spotted and Jefferson salamander accounts that hybrids may occasionally be tetraploid or higher ploidy and rarely are male, but this has been covered in the Ambystomatidae page that precedes them. There is no mention in the Red-backed Salamander account of the rare all-red (erythristic) phase without dark sides recorded from southern Ontario, an omission also imposed by space constraints in the single-page per species format. In accounts for Bullfrogs and Snapping Turtles the number of eggs is low, the for- mer can produce more than 8000 eggs, and the latter more than 40. Brown Snakes often have pink or white undersides as pointed out in the text of that species, but only light brown is used to distinguish it from the Red-bellied Snake in the text of the latter. The scientific name of the introduced Eastern Box Turtle is Terrapene carolina, not Terrapene ornata as given on page 162. Although the photographs are generally vivid and outstanding for their purpose, occasionally they are atypically light as in the top Mink Frog on page 93 (in comparison with northern populations I have seen) or too dark to show detail as in the center Mink Frog on page 93 (where the hind leg markings are thus obscured and look more band- ed than blotched, a Green Frog character). Another possibly atypical example is the center photograph of Butler's Garter Snake on page 135 which looks very like a Common Garter Snake from eastern Ontario in the greenish yellow of its stripe and labial (lip) scale colouration, the trace of black rather than The Amphibians and Reptiles of Alberta: THE CANADIAN FIELD-NATURALIST — ———_——— ie Theo 4 Vol. 116 - the broad chocolate margin beneath the lateral stripe, and the position of that stripe which does not seem to be significantly above scale row three except at the head. The Wood Frog pictures might usefully have included one with a mid-dorsal light line, a variant mentioned in the text. Northern Ontario populations can have quite high numbers of these striped individ- uals, a distinctive variant occuring only in this species among Canadian frogs. The Black Ratsnake pictures usefully could have included a blotched juvenile to contrast with the Milk Snake which occurs within its range (the Racer selections effec- tively do include a juvenile) and the Hognosed Snake selections could have included a black (melanistic) animal to contrast with the spotted phase shown. The very dark backgrounds used for tadpoles and salamander larvae are at first discon- certing but surprisingly effective (and the Grey Treefrog tadpole is outstanding). Some frog tadpoles are paler in colouration than I am used to in eastern Ontario, and it would have been useful for researchers to have had the locality of each specimen given for all photographs. The designers appear to have inverted two pictures (bottom of pages 107 and page 143). Such flaws are inconsequential in the overall effect and general quality of the book. It deserves, and certainly will have, very wide sales and longevi- ty. It is conveniently field-portable and has a place in every naturalist’s pocket from eastern Manitoba to Prince Edward Island as, aside from the Spring and Mountain Dusky salamanders of the mountains of southern Quebec and the marine turtles of the Atlantic, it covers every species to be found in east- ern Canada, and most of those in the northeastern United States. The author and the talented designers at the Royal Ontario Museum should be congratulat- ed not only on the quality of this effort, but also in continuing a tradition of one of the things that a sci- entific staff of museums can do best — authoritative, accurate field guides. FRANCIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada A Field Guide and Primer of Boreal Herpetology, Second Edition By Anthony P. Russell and Aaron M. Bauer. 2000. Uni- versity of Calgary Press, Calgary, Alberta. 280 pages, illus. $24.95. This is a first in Canadian herpetological field guides, a second edition, and only seven years after the first appeared in 1993 (see review by James D. Gardner. 1994. Canadian Field-Naturalist 108: 379-380), a fine tribute to its initial success as both a field guide and introductory herpetological text, validating the judgement passed by the earlier reviewer. It has the conventional species accounts of 18 doc- 2002 umented species [10 amphibians (2 salamanders and 8 frogs and toads) and 8 reptiles (1 turtle, 1 lizard, and 6 snakes)] and four additional accounts of species that might occur in the province (1 frog, 1 turtle and 2 snakes). These numbers are unchanged from the first edition but the nomenclature has been updated as five species have undergone scientific name changes (Scaphiopus bombifrons to Spea bombifrons, Pseudacris triseriata maculata to Pseudacris maculata, Rana pretiosa to Rana luteiventris; Phrynosoma douglassii brevirostre to Phrynosoma hernandesi, Pituophis melanoleucus to Pituophis catenifer, and a sixth (Ambystoma tigrinum) is “in a state of flux” (page 1x) and western populations have been designated Ambystoma mavortium but this has not been generally accepted, and is not followed here. Research has continued on the herpetofauna in the interval between editions, particularly as part of the production of status reports on the wildlife of Alberta, eight of which to publication had dealt with amphibians and reptiles. As well, research on species that occur in Alberta has continued elsewhere in their ranges. References directly relevant to the her- petofauna of Alberta have increased by more than 150, and the text has been updated throughout to reflect this. In the species accounts a section on Conservation Status has been added. Errors in the first edition have been corrected, for example the first appearance of reptiles (page 13) is now pushed back to the Carboniferous Period (about 330 million years ago) from the Permian (260 million years ago) BOOK REVIEWS 655 in the first edition. The spot distribution maps have been updated from the research for status reports and other publications since the previous edition. Particularly significant examples are additions of records of the Long-toed Salamander, Ambystoma macrodactylum, for the Peace River region and dele- tion of a perplexing record for the Tiger Salamander, Ambystoma tigrinum, north of 53°. Records for the Spadefoot Toad extend its range in eastern Alberta north to between 52 and 53° from 51°N. The Boreal Toad, Bufo boreas, has now been extended east at about 51°N, nearer to the Saskatchewan boundary. Other records expand the range of the Columbia Spotted Frog, Rana luteiventris, somewhat eastward and the Western Hognose Snake, Heterodon nasicus, marginally to the west. Some species, particularly the Great Plains Toad, Bufo cognatus, the Canadian Toad, Bufo hemiophrys, and the Leopard Frog, Rana pipiens, show many new localities within their previ- ously known ranges. The concluding overview text sections retain headings of zoogeography, natural history, coping with cold, the challenge of aridity, defense, and venoms. The concluding section, “Man and the herpetofauna” in the first edition, has been retitled “Amphibians, Reptiles, and People”. New para- graphs updating conservation concerns have been inserted. FRANIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada Rare Amphibians, Reptiles, and Mammals of British Columbia By Sydney G. Cannings, Leah R. Ramsay, David F. Fraser, and Mark A. Fraker. 1999. Lands and Parks, Wildlife Branch and Resource Inventory Branch, Ministry of Environment, Victoria, British Columbia. 190 pp., illus. This coil-bound publication is a valuable resource summary reference that covers all native amphibian, reptile, and mammal species, subspecies or distinct populations that are included in the Red and Blue lists of species considered at risk in British Columbia. The information is stated to have been subjected to extensive technical review, but only contributors of information, not reviewers, are listed. Statements in the accounts are referenced to pub- lished scientific literature. The Red List includes those for which there is most concern, any indige- nous species or subspecies extirpated, endangered, or threatened in British Columbia. The Blue List covers species or subspecies vulnerable in British Columbia, those judged particularly sensitive to human activities or natural events, and also species that are suspected of being vulnerable but for which current information is insufficient to designate them in another category. One declared use of the Red and Blue lists is “in the Forest Practices Code to identify endangered or threatened wildlife that require atten- tion under forest development, silvicultural, and range use operational plans”. An introduction which gives definitions used and ‘book's scope is followed by accounts for each taxon by group. Each section has its own reference list. Four appendices give the 1998 Red and Blue lists, the status of the listed taxa in adjacent jurisdictions, and the global and provincial ranking of vertebrates in British Columbia. The 73 taxa accounts, 146 pages, include: 8 amphibians (3 salamanders and 5 anurans), 10 rep- tiles (2 turtles, 1 lizard, 7 snakes); and 55 mammals (4 shrews, | mole, 8 bats, | pika, 2 hares, | rabbit, 2 mountain beavers, | marmot, | ground squirrel, 4 chipmunks, | pocket gopher, | pocket mouse, 656 3 voles, 1 lemming, | harvest mouse, 1 jumping mouse, 2 bears, | sea lion, 1 Sea Otter, 2 wolverines, 1 Fisher, 3 weasels, 1 badger, 3 ecotypes of the Killer Whale, 1 porpoise, 1 elk, 1 caribou, 2 bison, 3 sheep (Bighorn and Thinhorn). Each account gives paragraphs on Taxonomy, Status (Global, IUCN, COSEWIC, Provincial), Global Range, Ecoprovinces, Biogeoclimatic Zones, Biology, Global Ranking, Provincial Ranking (Rank and Comments), Range in British Columbia, Esti- mated occurrences, Abundance, Trend, Protected occurrences, Threats, and Conservation Needs (Research, Inventory, Management and Stewardship) and a map showing shaded distribution in British Columbia. The accounts are brief summary compila- tions from the literature, apparently with little new material. However, most statements have citations of source, allowing original studies and greater detail to be found. Most references refer to data from British Columbia populations. The usefulness of the maps is diminished, however, by failure to use dots showing documented localities. The evaluation of threats section ocasionally seems generalized from the entire range and simply extrapolated to British Columbia without justifica- tion, for example in the Pacific Pond Turtle. On page 29 it is stated for it that “The major threats are loss THE CANADIAN FIELD-NATURALIST [| Be Sse Vol. 116 “ of habitat through urban and agricultural develop- ment in the Lower Mainland, and the introduction of non-native predators.” A surprisingly focused assess- ment when there are no British Columbia specimens of this species and the only two documented obser- vations of it for the province, both in the 1930s, are from the city of Vancouver, with one additional vague undocumented “record” suposedly from 1966. Little useful purpose is served by even including a species on such evidence, let alone trying to guess threats. It was comonly marketed for food in its range to the south, and individuals reported were most likely imported escapes, rather than last survivors of a overstressed native population. No mention is made of two British Columbia turtle specimens collected in 1929 (Canadian Museum of Nature, 1525 from Alberni, and 1551 from Vancouver) of an Asiatic species, Chinemys reevesii, which clearly demonstrate the release or escape of turtles that were imported and marketed in both Victoria and Vancouver as is documented by Storer in Copeia 1937(1): 9-11. FRANCIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with Comments Regarding Confidence in Our Understanding By Committee on Standard English and Scientific Names, Brian I. Crother, Chair, Jeff Boundry, Jonathan A. Campbell, Kevin de Queiroz, Darrel R. Frost, Richard Highton, John B. Iverson, Peter A. Meylan, Tod W. Reeder, Michael E. Siedel, Jack W. Sites Jr, Travis W. Taggart, Stephen G. Tilley, and David B. Wake. 2000. Society for the Study of Amphibians and Reptiles, Herpetological Circular Number 29. 82 pp. [Available from SSAR Publications Secretary, Robert D. Aldridge, Department of Biology, Saint Louis University, St. Louis, Missouri 63103 USA; e-mail: ssar@slu.edu]. U.S. $10.00. The drive to standardize common names for North America amphibians and reptiles has been strong in herpetology for the past 50 years, the one to compile checklists of accepted scientific names has been longer, 85 years. Combined, they are still very much a work in progress. The process of compiling the accepted scientific names in a checklist reflecting the opinions of a cross-section of herpetologists rather than that of a single expert was begun in 1917 by Leonard Stejneger and Thomas Barbour. They updated their list four times in 1923, 1933, 1939 and 1943, but all omitted common names. Although multiple alternative common names usually listed in regional or single-group publica- tions, K. P. Schmidt in the 6th Checklist of Amphibians and Reptiles of North America, pub- lished by the American Society of Ichthyologists and Herpetologists (1953) may have been the first to attempt application of a single standardized com- mon name for each taxon for the entire North American herpetofauna. In the same year the American Society of Herpetologists stuck a commit- tee, chaired by Roger Conant, to revise common names, and produced a list published in Copeia 1956: 192-185 in time for use in Conant's pioneer- ing field guide to eastern amphibians and reptiles for the Peterson Series, as part of an expansion of the series beyond the initial pioneering guides in birds. In 1978, a committee chaired by J. T. Collins produced a revised list for the Society for the Study of Amphibians and Reptiles (SSAR Herpetological Circular 1: 1-36) and Collins went on to produce revisions in 1982 (SSAR Herpetological Circular 12: 1-28), 1990 (SSAR Circular 19: 1-41), 1997 (SSAR Circular 25: 1-40). The latest checklist, by a new committee, is a marked improvement over all previous editions as 2002 for the first time changes in scientific names from the last edition are not only referenced to recent liter- ature but conflicting opinions discussed. Now read- ers can readily track and evaluate decisions for themselves. Groups have been split among individu- als or subcommittees: frogs: Frost; salamanders: Highton, Tilley (Chair), Wake; lizards: de Queiroz (Chair), Reeder, Sites; snakes: Boundry, Campbell, Crother (Chair), Taggert. There is also a discussion of new rules applied in the formation of common names to provide more consistency, largely following the lead set by the American Ornithologists Union, which pioneered standardization of common names for birds long before the herpetologists attempted it. Numerous changes have been invoked. Just as we were getting finally accustomed to using Redbelly Snake, and similar contractions (Ringneck Snake, Redback Salamander, Hognose Snake, etc.) for compound names where a character was singular in an individu- al, we find the new list reverts to the traditional Red- bellied Snake, etc., a form abandoned earlier as suggestive of an individual having multiple bellies. For a character with several occurrences in one indi- vidual, such as in Blue-spotted Salamander, the name is unchanged but the distinction between sin- gular and plural states is now lost. Many other com- pounds have been changed to provide consistency. Thus the familiar Garter Snake becomes Gartersnake and Rat Snake becomes Ratsnake, presumably because the long used Rattlesnake has become the model. Other reversions invoked here are the undig- nified Stinkpot back again for Sternotherus odoratus. This had been changed to Common Musk Turtle in recent lists but the present compilers were apprehensive that “Common” might be misinterpret- ed to imply abundance rather than widespread! Problems arise in compiling common names for both species and subspecies. Confusingly, some- times completely unrelated names are still used for a species and a contained subspecies, such as Notophthalmus viridescens being designated the Eastern Newt, where as its nominate subspecies Notophthalmus viridescens viridescens is the Red- spotted Newt. [t seems unfortunate that this commit- tee has found it necessary to add suffixes to clarify (for those unfamiliar with amphibians and reptiles?) long recognized names such as Leatherback which becomes Leatherback Seaturtle. Fortunately atroci- ties like “Racer Snake” favoured by some Canadian conservationists apparently faced with considering reptiles for the first time, are not invoked. (Who would impose, for example, “Oriole Bird” to satisfy those unfamiliar with avian lifeforms?) Regrettably, some Canadian clarifications I proposed in 1984 (Introduction to Canadian Amphibians and Reptiles, National Museum of Canada), and followed for a time by some Canadian workers, such as Yellow- BOOK REVIEWS 657 spotted Salamander for Ambystoma maculatum, to distinguish it from the Blue-spotted Salamander; Midland Chorus Frog for Pseudacris triseriata which is not a “western” species in Canada; or the distinguishing and informative names Tetraploid and Diploid Grey Treefrog for Hyla versicolor and H. chrysoscelis, continue to be rejected or ignored here as in the previous recent American lists. Although subspecies status for the Canadian Toad as Bufo americanus hemiophrys is still not accepted for this list, at least a reference to it as a possible philosophi- cal treatment is included. A positive innovation, instigated for the first time in a sanctioned list for amphibians and reptiles, is the capitalization of the first letter of each word in all common names of species and subspecies, to treat them as proper nouns, a practice borrowed from the American Ornithologists Union checklists for birds (and followed for 20 years in The Canadian Field- Naturalist for species names of all animals and plants). This new checklist will be widely welcomed and followed. It is particularly valuable for consitancy of use by popular writers and conservation agencies (and editors) as it gives them an updated source of standard usage. However, it is a myth that common names are stable; every list sees some changes even of frequently used names. Unlike changes in scientif- ic names which reflect new knowlege of relation- ships which necessitate change, those in common names seem to change on whim mainly with changes in committee membership. It is important especially for the non-herpetologist to note, however, that use of any list is not mandatory, it is simply the consen- sus opinion advanced by one expert committee at the time, and as such it is advisory only. Scientifically, no list can do more than represent the state of the art at one point in time, and becomes outdated at the rate of new work that is constantly in progress. Since the appearance of this list, proposals have been published to remove the Wood Turtle from the genus Clemmys and resurrect either Glyptemys (Holman and Fritz, 2001, Zoologische Abhandlungen, Museum Tierkunde Dresden 51(20): 331-354) or Calemys (Feldman and Parham, 2002, Molecular Phylogenetics and Evolution 22(3): 388- 398) from the synonymy (both first used in Agassiz, 1857, Contributions to the Natural History of the United States of America) for it and the extra- Canadian Muhlenberg's Turtle. Holeman and Fritz would apprear to have publication priority as first revisors. Both papers also suggest that the Pacific Pond Turtle should also be removed from Clemmys (leaving the latter monotypic with the Spotted Turtle, Clemmys guttata, as its sole species) and transferred with Blanding's Turtle into subgenera (placed in parenthesis) into the genus Emys as Emys (Actinemys) marmorata and Emys (Emydoidea) 658 blandingii. An alternative is to give both monotypic genus status and Actinemys marmorata and (as presently) Emydoidea blandingii. Another change since the checklist is the separa- tion of the Prairie Rattlesnake and North Pacific Rattlesnakes as Crotalus viridis viridis and Crotalus oreganus oreganus (Ashton and de Queiroz, 2001, THE CANADIAN FIELD-NATURALIST OO ———_ TLE ea | Vol. 116 - Molecular Systematics and Evolution 21(2): 176- 189). FRANCIS R. Cook Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada Pigeons and Doves: A Guide to the Pigeons and Doves of the World By David Gibbs, Eustace Barnes, and John Cox. 2001. Yale University Press, New Haven and London. 615 pp., illus. U.S. $60. This is another addition to the collection of books on bird families and groups that have been appearing over the last 15 or so years, perhaps triggered by the success of Harrison’s 1983 volume on seabirds. All share an overall similarity in style, which will be familiar by now to most readers interested in this field, but vary widely in their approach to the details of their accounts. Harrison’s book filled an important need for birders in providing an authoritative identi- fication guide to a group of birds only poorly covered by other works. Some subsequent books in the genre have also emphasized identification, often with less success, while other authors have appeared to concentrate more on broader systematic and life history details. The present volume deals only briefly [under three pages] with the characters of the family as a whole, and is no more than adequate in its coverage of identification. Its strength lies in the species’ accounts, which provide a good summary of details relating to each species. Each occupies about a page and a half, plus a range map, and covers identification, voice, habits, habitat, and status and distribution, followed by a detailed description, with key measurements, and an outline of geographical variation. There is real merit in this approach, as the Handbook of the Birds of the World, Volume 4 [Del Hoyo et al. 1997], pro- vides a most extensive and lavishly illustrated account of the family as a whole, but only relatively terse sum- maries for each species. By the same token, identifica- tion problems are well covered in field guides for most of the more popular birding destinations. Looking at the species’ accounts in more detail, I found that in general the detailed coverage seemed quite good, although noticeably weaker on North American topics. The range maps are erroneous for several species: for example, Mourning Doves are shown as non-breeding visitors in Canada and the northern US. The text is more accurate, but still implies that the Mourning Dove occurs no further east in Canada than southern Ontario. Similarly, nei- ther text nor map shows White-winged Dove occur- ring in Florida. Curiously, the “Similar Species” entries for both the above birds discuss Eurasian Collared-Dove, but the text on the latter makes no mention of it occurring in North America at all! One of the problems facing birders here when identi- fying this species is separating it from the domesti- cated Ringed Turtle-Dove, usually referred to as Streptopelia risoria. This bird occurs quite frequent- ly in the wild as an escapee, and there are well-estab- lished local populations of feral birds. Both the identification and systematics aspects of this subject have been well discussed in the literature, but all of this is wholly ignored in the current volume. The colour plates follow the usual pattern, grouped at the beginning, with the images on the right and text on the facing page. The latter gives a line on distribution, and identification criteria for each image. Usually between 4 and 6 species appear on each plate, and as many as 19 separate images. Consequently some of the plates appear rather clut- tered, but never confusingly so. The paintings are all by two of the authors, Barnes and Cox. Some are more convincing than others: I found the Australian bronzewings very disappointing, and the normally elegant Crested Pigeon looks downright ill. As a whole, the Handbook of the Birds of the World plates are more satisfying [Cox contributed to both], although I liked the small flight portraits in this work. There are no images showing the plumage variation in feral Rock Doves, although there is an entire plate devoted to the Passenger Pigeon. Overall I found Pigeons and Doves rather disap- pointing. The errors in distribution information leave a sense of uncertainty about the text as a whole, identification details are often more thoroughly coy- ered elsewhere, and the quality of the plates is uneven. The book also suffers from the problems of some others of this genre, as its intended audience seems rather ambiguous. If your interest is in the Columbidae, then this book could provide a useful compendium of information on the species in the family, but I doubt if the more general reader will find it worth the price. CLIVE E. GOODWIN 1 Queen Street, Suite 401, Cobourg, Ontario K9A 1M8 Canada 2002 BOOK REVIEWS 659 Handbook of Birds of the World Volume 7 Jacamars to Woodpeckers Edited by Josep del Hoyo, Andrew Elliott and Jordi Sargatal. 2002. Lynx Edicions, Barcelona, Spain. 613 pp., illus. U.S. $185. I thought the arrival of Handbook of Birds of the World Volume 7 would mark a major milestone — the authors would be half way though their epic task. This is no longer true. After broad, worldwide con- sultation they have decided to publish 16 volumes to accommodate all the detailed material available. I have not seen a final publication date for this marathon task but it will likely be a dozen or more years away. In fact the elapsed time so far has been long enough for the Atitlan Grebe to go from “almost extinct “(Volume 1) to “extinct” (Volume 7). The change to 16 volumes, at about $200 a volume, will also add $800 to the total price. The authors have at least completed the non-passerines and to mark this point they have issued a plastic insert. This single sheet lists the families and genera in volumes | to 7 and gives volume and page numbers for each genus. Handbook of Birds of the World Volume 7 opens with an essay on 132 extinct species. That is those species for which there is absolutely no hope of being found alive again. (Species for which there is hope, however small, are contained in the main text. This means Ivory-billed Woodpecker is kept as an extant species based on the recent Louisiana report.) Also discussed are the mystery birds — birds that have been described but for which there is no hard evidence of their existence. This is a fas- cinating account of birds well-known and obscure that come from a shadowy world of science blend- ed with half truths, mistakes and over zealousness. It provides a fine background to extinction and sets the stage for understanding the fragility of some currently endangered species. I was particularly interested in the author’s comments on the Tachira Emerald (Polyerata distans). First, I found the index was in error by a page. The authors referred to this bird as a probable hybrid (presumably Glittering-throated Emerald [Polyerata fimbriata] x White-chinned Sapphire [Hylocharis cyanis]) but gave little discussion except to suggest it was prob- ably not extinct. This volume covers jacamars to woodpeckers, in all just over 400 species. With all of the species cov- ered it is easy to place them in their family. All the jacamars, for example, are similar and typically are chestnut and emerald. Likewise the puffbirds can be characterized as brown, black, and dumpy. The woodpeckers, which constitute over 45% of this vol- ume and account for 55% of the species, fall into characteristic groups. All are immediately identifi- able as a woodpecker, and then it is simple to clump them into distinctive groups. For example, the flick- ers are a very obvious coherent unit. It is not until you look at the barbets and the toucans that you see some wild diversity. Even here it is not the body shape but the colour that varies. The multi-coloured toucans and barbets are South American. The more uniform brown species of barbets are African and the green ones are Asian. | did note that the Gilded Barbet, Capito auratus, (a very nice depiction in the plates) is lumped as the Black-spotted Barbet, Capito niger auratus. Other recent publications have given this bird full species status. The section covering each family begins with a summary box which gives basic information on clas- sification and range. It shows silhouettes of the species within the family compared to that of a human. This summary provides a quick visual intro- duction to the main text. This text for the introduction of the family is reasoned, current, and instructive, and it provides a solid basis for understanding the family while pro- viding a point of reference for future studies. The content of the individual species descriptions is informative, providing a satisfactory level of essential information while avoiding cloying detail. The book ends with about 4000 bibliographical references. It is interesting to note that DNA studies are hav- ing a significant impact on our ideas about species. In this volume the authors split the European and American Three-toed Woodpeckers based on DNA results. They further suggest that other disjunct populations may also be separate species. The artwork lives up to the typical, high HBW standard. I scanned through looking for errors and found none, just a few minor discrepancies in depth of colour. Most notably the wings on the Cream- coloured Woodpecker are not brown enough. This is actually confirmed by a photograph of this species earlier in the text. Additionally, | noted there are more representations of females than in earlier edi- tions. The over 300 colour photographs run from good to great. The photographs are not merely an extension of the plates. Rather they are a comple- ment to the text and plates. They highlight aspects of the bird’s biology and behaviour, coupled with a glimpse of its habitat, leaving the artwork to provide a means of identification The range maps are clear and show the currently accepted distributions. Whether they are accurate Is a little more difficult to judge. Many of the species live in remote areas that are not well known. The map size 5 X 3'/2 cm is quite small. When com- pared to Birds of the Western Palearctic’ or Birds of Canada? this map size is a little difficult to interpret. For example, it is not easy to see the difference in range between the Emerald Toucanet and the Chestnut-rumped Toucanet which, in reality, are sep- arated by the Andes mountain range. 660 So this great contribution to ornithological litera- ture steam rollers on slowly transforming itself into the bible of birdwatching. It is hard to believe that I will be in my seventies before the last volume is pro- duced and the creators will then have to consider revising Volume 1. References IThe Birds of the Western Palearctic, Chief Editor: the late Stanley Cramp, Editor of Vols. I-IV: K. E. L. Simmons, and THE CANADIAN FIELD-NATURALIST Vol. 116 - Editor of Vols. VII-IX: C. M. Perrins, 1994. Oxford University Press, England. ?The Birds of Canada, Bulletin 203 (Biological Series 73), W. Earl. Godfrey, 1966. National Museum of Canada, Ottawa, Ontario. Revised Edition 1986. Roy JOHN 2193 Emard Crescent, Beacon. Hill North, Ottawa, Ontario K1J 6K5 Canada The Marbled Murrelets of the Caren Range and Middlepoint Bight By P.H. Jones. 2001. A Western Canada Wilderness Committee Research Report. ISBN 1-895123-13-5. 150 pp. $34.95. “British Columbia is endowed with one of the most diverse and probably the richest natural her- itage in Canada and yet so little is known of this vast resource”. This book is not strictly a scientific one, but surprisingly, the first of its kind. It provides the reader with an alternative and widely shared view on the Marbled Murrelet issue in southern British Columbia. It is about the classical conserva- tion battle for a habitat and a species of world importance: the Marbled Murrelet (MaMu), Brachyramphus marmoratus, and its nesting habitat, the coastal Old-Growth Forest. The author states “I was alarmed at what was happening on the Caren and elsewhere in the province. My respect for the profession of forestry had clearly suffered a huge shock’. The book is for MaMu-holics written by a Murrelet-phil. But it is also suggested reading for any naturalist and wildlife ecologist in the Pacific Northwest and beyond. Readers learn from a first-hand account on “how to find a Marbled Murrelet nest’, as well as about many aspects of the fascinating biology of this bird, including all the politics around it. Certainly, the finding of the first active Marbled Murrelet nest in Canada on 12 August 1993 makes for a central theme of this book. The discovery pre- sented a major effort over many years, based on a tremendous amount of amateur field-work and vision; eventually it turned into a success. However, the events that led to this historical finding deserve some attention. For instance, it is not clear why the people who first located the nest were not authors in the forthcoming publication. The overall motivation and funding situation for “The Caren” remains somewhat unclear as well. Many fascinating details about this bird, its habitat (including the sea) and its inhabitants can be learned from this publication, too. For instance, the Caren Range claims the oldest tree in Canada with a con- firmed age of over 1700 years; the discovery of this finding is worth a read in itself. Well-rounded, the text even presents “MaMu-history” and native aspects. And certainly, a fair bit of lifestyle in coastal British Columbia is described. The Marbled Murrelet is not only a species difficult to study, but also to draw. The author includes tasteful and nice black-and-white illustrations of this bird. The statement of the author that Marbled Murrelets are unlikely migratory can be confirmed (wintering birds are found consistently along the British Columbia coast). Although not well liked by Canadian governments, this fact has major conserva- tion implications because according to the Bird Migratory Act 1917 the federal government can only support migratory species; this bird not being migra- tory would drive the responsiblity in the hands of the provincial government. The author states on this sub- ject “...Provincial authorities in British Columbia have done virtually nothing to protect marbled mur- relets and to set them on a path of recovery’. It is worthwhile to mention that besides facilitation, the federal goverment has also not provided much guid- ance in the last years on this matter. Several interesting biological features about Marbled Murrelets are reported in this book: loca- tions and timing of moult. Other jewels in this book might be the raw data on nest visits, and pair-bond- ing described for October. The described mating observations, the murrelet foraging process, and transformations of summer and winter plumages in September are also precious. The claim that Marbled Murrelets could be identified individually, and that males could be distinguished from females in the field, deserves more proof; at least this has never been reported anywhere else in the world. Another concern is that Mew Gulls don't depend on the Old- Growth Forest, and re-use of Marbled Murrelet nests is seen as a rather rare event, so far. I would doubt, too, that this bird is colonial, as hinted in the text. Unfortunately, the statements about the Simon Fraser University are disagreeable and need correc- tion (e.g., a cliffnest was indeed found by this group, and over 150 nests were located in the last four years 2002 with radio-telemetry from helicopter). For these subjects, the reader is suggested to contact the researchers from the Centre for Wildlife Ecology directly, or check their website (URL: http://www. sfu.ca/biology/wildberg/species/mamu.html). Besides the claim of the author and the obvious loss of Old-Growth Forests, there are still no statistical indications that the populations in British Columbia overall have in fact declined — a topic of major con- troversy. A less disputed point is made in this book that the traditional hype about the nesting habitat is not really reflecting the overall life-needs of the Marbled Murrelet; the marine aspects of this bird are even less studied, nor do any (!) Marine Protected Areas exist along the unique coastline of western Canada (Middlepoint Bight as a feeding area might present one candidate). The issue of the effect corporate funding would have on science, and on Marbled Murrelet research in particular, is still open for discussion, but is heavi- ly argued, here. Since controversial topics are report- ed on, names like Randy Stoltman, Mike Harcourt, and Glen Clarke are not missing in this book. The footprints left by prominent Marbled Murrelet researchers in British Columbia like Spencer Sealy, Harry Carter, Jean-Pierre Savard, Gary Kaiser, and others are found throughout. “We made our point, too, with the Forest Service that it was they who were forcing the companies to get out there and log the Caren, even when there was a government process in place to protect areas like the Caren!”. Obviously, the management of the forests in British Columbia gets much debated in this book, as elsewhere. “We cannot have a Ministry of Environment which is subservient to a Ministry of Forestry as has been the case in the past”. The term “overcutting” is mentioned many times by the author. It is not clear how this can occur in a country that claims to do “sustainable forestry”; apparently without any relevant and consistent ground-based inventory of Old-Growth Forests! Jobs matter in British Columbia, but due to odd and failed manage- ment, the international forest industry — to name Japan, USA, and even Norway — has become the main drivers of the agenda of forests in coastal B.C. BOOK REVIEWS 661 The author refers often to the “Marbled Murrelet Recovery Team”, which is supposed to do something good for the Murrelets. The Protected Areas Strategy (PSA) and the BC Forest Practices Act might be one approach. However, other solutions exist, and as this book clearly indicates, nests are not found in low elevation but rather at higher elevations (>400 m) and with steeper slopes. The least one could say about all this is that Marbled Murrelets are very robust in their nesting habitat, which covers appar- ently a wide range of landscape and tree characteris- tics. This puts doubt on the Marbeled Murrelet as an indicator species, since it “indicates” so many habitat features (way beyond the low elevation forests, a tra- ditionally claimed prime nesting habitat for this bird). Biases in Marbled Murrelet research and con- servation are introduced by investigation approaches with a poor research design, by the management term “forest stand” (basically impossible to define objectively in the field and from maps), by using the term Old-Growth Forest (over 12 definitions exist), and by the current counting scheme for Murrelets in the forest (titled “Occupied Detection” surveys). As the author confirms, the Canadian counting protocols used for Occupied Detections and at-sea surveys dif- fer markedly from the ones used in the United States. The author is certainly not an “ecoterrorist”; instead, he is a devoted naturalist and deeply con- cerned about his “backyard”, the Caren Range and Middlepoint Bight. His book provides an emotional and passionate narrative. It presents a textbook- example to concerned citizens and naturalists about his battle with local experts, the local District Forester, and governments to convince and to achieve his case; these are experiences one makes when interacting in “territories” of the established and so-called local experts. This recommended book shows how he succeeded. FALK HUETTMANN Geography Department, Earth Science, 2500 University Drive N.W., University of Calgary, Calgary, Alberta T2N 1N4 Hummingbirds of North America: The Photographic Guide By Steve N. G. Howell. 2002. AP Natural World. Academic Press, San Diego. 222 pp,. illus. U.S. $24.95. Photographic bird guides have a rather chequered history. They seem like a good idea, but in practice they have all too often proved to be less than ideal, with poor poses, ambiguous colours, and a lack of uniformity all combining to reduce their utility. So I approached such a guide to hummingbirds with con- siderable skepticism after all, hummingbird colours can be difficult to determine at the best of times, and their behaviour doesn’t help much. But Steve Howell has surmounted all the usual problems. This is the ideal photographic guide, with a multitude of carefully selected poses, detailed information in the captions, and meticulous attention to the appearance of each image. 662 It has a text to match. The highlights of the 34- page introduction are a section entitled “taxonomy and identification framework”, which includes a review of the characters of the North American gen- era, and a very detailed and comprehensive outline of field identification. The latter — like the rest of the book — has large and carefully annotated illus- trations which complement the text very well. The author’s own words summarize the challenge this barrage of information poses to the reader. He says: “While the foregoing may seem an almost over- whelming amount to digest, there is no hurry”. This section does indeed put a whole new perspective on hummingbird identification, and presents a whole new set of challenges to those of us in the east. So I greatly appreciated his comment. It neatly sums up his practical, down-to-earth approach to the entire book. The heart of the guide is a detailed account of the identification of the 22 species of hummingbird that either breed or have occurred in North America north of Mexico, plus Cuban Emerald and Bumblebee Hummingbird, for which reliable sight records or specimens exist. Treatments range from some 12 pages each [including four devoted to 11 or 12 photographs] for widespread and tricky species such as Ruby-throated and Black-chinned, to four THE CANADIAN FIELD-NATURALIST ee cr eee ge ee a ec Vol. 116 - pages for easier and less common species. In fact, the coverage of Ruby-throated and its “look-alikes” also benefits from a five-page discussion on the “small gorgeted hummingbirds”. All the species are dealt with genus by genus, and have a short general statement on the characters of the genus as a whole at the beginning of each section. The book concludes with a glossary, a list of references, and a short index. This is an excellent, carefully conceived, well- written and practical guide, obviously by someone who has an intimate familiarity with his subject. I found the occasional mistake, and the approach to reference citations was a bit confusing at first, but overall it is hard to find fault with this book. It’s yet another in the sequence of superb guides that have been published in the last few years, and it’s one that will prove invaluable to easterners like myself who suddenly find themselves faced with an array of these enchanting yet pernickety little creatures on a trip out west. I think I’ll go back to Arizona and try it. CLIVE E. GOODWIN 1 Queen Street, Suite 401, Cobourg, Ontario K9A 1M8 Canada State and Provincial Amphibian and Reptile Publications for the United States and Canada By John J. Moriarty and Aaron M. Bauer. 2000. Society for the Study of Amphibians and Reptiles, Herpetological Circular Number 28. 52 pp. [Available from SSAR Publications Secretary, Robert D. Aldridge, Department of Biology, Saint Louis University, St. Louis, Missouri 63103 USA; e-mail: ssar@slu.edu]. U.S.$8.00. This bibliography compiles North American lists and guides for 51 states of the U.S. and 10 provinces and three territories (plus a general category for the Canadian Maritimes). The earliest entries are Storer 1840 for Massachusetts and DeKay 1842 for New York. In all, 835 entries are listed, or 881 if multiple entries under several categories are individually counted. In the United States Florida leads with 46 citations; in Canada, British Columbia has the most, 10. The scope is limited to publications dealing with a single state or a significant portion of one. Publications dealing with more than three states or provinces have been excluded, except for the historic regional units of New England (Connecticut, Maine, Massachusetts, New Hampshire, Rhode Island, and Vermont) and the Maritimes (here defined in error as New Brunswick, Newfoundland, Nova Scotia, and Prince Edward Island - in proper eastern Canadian usage the four together are called the “Atlantic provinces” and “Maritimes” is restricted to refer- ences to the three that joined Canada in the 1800s, excluding Newfoundland which did not join until 1949). This publication serves as a handy reference for entrance to provincial and state literature and should have wide circulation and use. Of interest is that seven of the Canadian entries are to papers in The Canadian Field-Naturalist and its predecessor The Ottawa Naturalist. A major historic omission for Ontario is J. H. Garnier. 1881. List of Reptilia of Ontario. Canadian Sportsman and Naturalist (Montreal) 1(5):37-39 and, for Quebec, Leon Provancher 1874. Faune Canadien. Les reptiles. Le Naturaliste canadien 6: 273-278; 289-298; 321-330; 353-370. and 1875. Faune Canadien. Les reptiles. Le Naturaliste canadien 7:10-20;42-46:65-73. Its usefulness could have be enhanced consider- ably by including a section for the major regional guides to larger units in North America both for amphibians and reptiles, and for the groups within them. FRANCIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 2002 Salamanders of the United States and Canada By James W. Petranka. 1998. Smithsonian Institution Press, Washington and London. xvi + 587 pp., illus. + plates. U.S. $60. James Petranka, Biology Department of the University of North Carolina, has 20 years in the field as evidenced by 22 entries as senior author, the first in 1979, in the bibliography of this book. Among them are fieldwork oriented papers on winter effects on woodland salamander (Plethodon) popula- tions in Kentucky, effects of timber harvesting on southern Appalachian salamanders, and ecology of the Small-mouthed Ambystoma texanum and Marbled, A. opacum, salamanders. For this book he has set himself a daunting task. The last comprehen- sive compilation of North American salamanders was by Sherman C. Bishop (1943) in the Comstock series of classic “handbooks” pioneered by A. H. and A. A. Wright (who wrote the frog and snake vol- umes) from Cornell University. A new synthesis is especially long overdue considering the importance of the area covered. North America, with 9 families and 24 genera, has the greatest diversity of salaman- ders in the world. As well, from 1943 to 1997 more than 1500 refereed research papers on systematics, ecology and natural history of North American sala- manders have appeared and more than 30 new species have been described. Setting out his approach in his Preface, Petranka first faces the critical questions of taxonomy (nam- ing units) and systematics (arranging these units in relation to one another), both highly contentious top- ics among current salamander specialists who have varying criteria and philosophies (e.g., biological, evolutionary, and phylogenetic species definitions) guiding them. As a result, the number of species rec- ognized can vary from fewer than 110 to more than 150. Petranka recognizes 127, reflecting a moderate- ly conservative approach. The Preface is followed by an Introduction which gives brief overviews of content, diversity, ecologi- cal roles, life history, and general natural history. A section on Salamander Identification and Plan of the Book is next, then one on Conservation Biology of Amphibians, and Key to Adult Salamanders and Key to Larval Salamanders of the United States and Canada. The book concludes with a Glossary of 51 terms and a Literature Cited of about 2100 refer- BOOK REVIEWS 663 ences, Collection Localities (to county generally) and Photographic Credits for Color Plates, and a five-page index to genera and species. The bulk of the book (pages 35-492) is the family and species accounts. The former cover characteri- zation, fossil history, and major systematic treat- ments. The latter give scientific and common name and include sections on Identification, Systematics and Geographic Variation, Distribution and Adult Habitat, Breeding and Courtship, Reproductive Strategy, Aquatic Ecology (where applicable), Terrestrial Ecology (where applicable), Predators and Defense, Community Ecology, Conservation Biology, and (sometimes) Comments. The text of these sections is rich in references and emphasizes comparative geographic coverage of ecology, behaviour and life history. Within the species accounts there are one-column black-and-white photographs of adults (occasionally several views), and often of larvae, eggs, and spermatophores. There is a page-width continental map of the United States showing distribution as grey blocks for monotypic species and in varied hatching and marbling for subspecies of polytypic forms. For species which range north into Canada most provinces are added but the state of Alaska (Taricha granulosa) and the Labrador portion of Newfoundland (Eurycea bislineata, Ambystoma laterale) appear only where there are species rang- ing there. Finally, inserted in the center of the book are 172 colour photographs of adults and larvae, with subspecies included for a few polytypic forms (Ambystoma tigrinum, Notophthalmus viridescens, Eurycea longicauda, Gyrinophilus porphyriticus, Pseudotriton ruber, Taricha granulosa, Siren inter- media). This thorough and comprehensive book is a worthy successor to the pioneering text of Sherman Bishop nearly 60 years ago. It will be on the essen- tial reference shelf of every professional herpetolo- gist, and consulted for detailed information and references by a legion of field naturalists and conser- vation environmentalists of every hue. FRANCIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 664 THE CANADIAN FIELD-NATURALIST Vol. 116 ~ Patterns of Distribution of Amphibians: A Global Perspective Edited by William E. Duellman. 1999. The John Hopkins University Press, Baltimore and London. viii + 633 pp., illus. U.S. $69.95 The phenomenal growth of research on amphib- ians and reptiles in the last half of the 20th century, including the description of a multitude of new species to bring the world total to over 4700, ever more precise data on ranges, and fine-tuning of rela- tionships partly though the ever expanding use of DNA analysis, has made possible the new synthesis attempted here of the mosaic of individual ranges throughout the world. The first chapter is an overview, “Global Distribution of Amphibians: Patterns, Conservation, and Future Challenges” (William E. Duellman). This is followed by nine individually authored chapters on amphibian distri- bution divided geographically: “Nearctic Region of North America” (W. E. Duellman and Samuel S. Sweet) “Middle America” (Jonathan A. Campbell), “West Indies” (S. Blair Hedges), “South America” (W. E. Duellman), “North Africa”, “Europe”, “Western Asia’, and “former Soviet Union” (Leo Borkin), “temperate Eastern Asia” (ZhaoEr-Mi), “Southern Asia and adjacent islands” (Robert F. Inger), “Sub-Saharan Africa’, “Madagascar, and Seychelles” (J. C. Poynton), and “Australo-Papuan region” (Michael J. Tyler). Some of the contents were originally prepared for presentation in a sym- posium at the Second World Congress of Herpetology in Adelaide, Australia, in December 1993 but new data have considerably augmented the presentations. Duellman's initial overview chapter is comprehen- sively illustrated with graphs and tables of continen- tal comparisons and contrasts in number of families, genera and species, the rate of description of new species, as well as the geographic occurrence of endemism, all of which will be useful not just to every herpetologist but also to the non-herpetologist for contrast with their own groups, and to the general naturalist for perspective of where this group fits into the natural world. The discussions of conservation and the declines and the need for more research cover much that is familiar, but which can never be restated too often. Although the southern areas of the world have the major share of extant species (South America leading, with 2465), and much of the diversity of families and genera, it is the northern continents that will be of immediate interest to Canadian herpetolo- gists, ecologists, and naturalists, North America with 241 species, and Eurasia with 192. The Nearctic Region as defined by Bill Duellman and Sam Sweet (pages 11-110), has 90 anurans (frogs, toads, and kin) and 151 salamanders. Maps and discussions of a selection of generalized envi- ronmental divisions and specific variables are pre- sented. Major physiographic provinces are defined as (from west to east) North American Cordillera (Pacific Ranges, Intermontane Plateaus, Rocky Mountain System), Interior Plains (Great Plains and Interior Lowlands), Laurentian Shield, and Appalachian Highlands. The majority of species occur outside the maximum former extent of Wisconsin glaciation and therefore much of the dis- cussion is of patterns south of it. There are gener- alised maps of limits for the 15 most northerly species but the boundaries shown are unfortunately inaccurate in places. Surprising is the statement (page 83) that “there are no clear correlations of dis- tributional limits with temperatures”. The pioneering herpetofaunal zones defined for eastern Canada by J. S. Bleakney in 1958 (National Museum of Canada Bulletin 155) are not discussed nor is his paper even cited. My 1984 Introduction to Canadian Amphibians and Reptiles is, but the only other Canadian references included are Anthony Russell and Aaron Bauer's guide to Alberta species, two publications by David Green, his students and col- leagues, and one in which the master of freeze toler- ance, Ken Storey, participated. The northern Palaearctic with 69 salamanders and 106 anurans is covered by Leo Borkin (pages 329- 420) who effectively presents data debunking the concept of a Holarctic unit for amphibians. The latter has sometimes been invoked by past workers because of well-established faunal similarities at a species or subspecies level in some other terrestrial vertebrate groups (birds and mammals) and some invertebrate and plant groups. In contrast, amphib- ians (like reptiles) share no species or sister species between Old and New worlds. Only four genera of amphibians (one salamander (Hydromantes) and one each of frogs (Rana), treefrogs (Hyla), and toads (Bufo) are represented in both, and none of three frog genera are exclusive to the northern regions. Even on the family level, of the 5 salamander and 8 anuran ones in the Palaearctic and 9 and 8 in the Nearctic, only four of 10 salamander and five of 11 anuran (frogs and kin) that occur in the two regions combined are common to both. The book is an instant classic reference, rich in data and comparison and contrast, a tribute to the industry of all its authors, but especially of its editor who has been a ever-growing force in amphibian systematics and zoogeography over four decades. FRANCIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada 2002 The Amphibians of the Former Soviet Union By S.L. Kuzmin. 1999. Pensoft, Sofia, Bulgaria, and Moscow, Russia. vi + 538 pp., illus. U.S. $98 + U.S. $12 shipping. For Canadians, this fine book gives a view of essentially a faunistic “alternate universe” to what they are familiar with — an amphibian assemblage evolving independently from that of North America since the Miocene yet coping with the same northern environmental problems. Previous overviews have been available in English translation of Russian clas- sics textbooks; this is a new text written in English. The Preface comments on past major texts for the area, the former USSR retained here as the area cov- ered with the terse comment that “The Soviet Union is no longer respected as a single political entity because of its dissolution and the creation of the Commonwealth of Independent States (CIS) and several Baltic Republics”. The information presented is selected from the author's database of 4000 biblio- graphic sources [a vast majority not available in English] and 7500 localities; still in progress, expla- nations with diagrammatic illustrations of standard measurements used for various groups and life stages, and use of a compilation on ecology for each species. Chapters 1-5 (the first 99 pages) cover History, Faunistics and geography, Ecology, Anthropogenic pressure and conservation, and Keys for species identification. The book ends with a 66-page bibliography, 26 pages of detailed spot dis- tribution maps, 32 pages of habitat and species colour photographs, a 4-page glossary, 5-page sys- tematic index of amphibians, 4-page Subject Index, and finally a 3-page list of species and subspecies of the area covered. The bulk of the book, 300 pages, is Chapter 6, “Species accounts”. Genera are briefly characterized here as well, before each group of species they con- tain. Each species account is headed by English and Latin name, synonymy, and Russian, German and French names. The sections for most include Taxonomic notes, Description (Morphology and Karyology), Distribution, Subspecific differentiation, and Variability, Ecology (Habitats and abundance, Thermobiology and Activity cycles, Reproduction, Development, Feeding, Natural enemies, parasites and diseases), Influence of anthropogenic factors, Status and conservation, and References. In all, 40 species (27 frogs [28 if the “klepton” Rana esculenta is included] and 13 salamanders) are represented compared to 44 (24 frogs and 20 sala- manders) in Canada. At the family level the Pelobatidae is represented by the genus Pelobates (2 species) instead of the North American endemics Scaphiopus and Spea) whereas Bufonidae, Hylidae and Ranidae have genera in common, Bufo (7 species), Hyla (3), Rana (1 1). The toads are in two species complexes: the Common Toads, Bufo BOOK REVIEWS 665 bufo (3 species) and the Green Toads Bufo viridis (4). Both groups lack cranial crests in common with the North American Western Toad, Bufo boreas. All the hylids are the European Treefrog, Hyla arborea, complex, and look superficially similar to the North American Pacific Treefrog, Hyla regilla [now included in Pseudacris by most authors]. In the ranids, the brown frog group has some species that resemble the North American Wood Frogs, Rana sylvatica, in having dark snout “masks” and some having dorsal stripes. But many have red ventral colouration, not seen in Wood Frogs but present in some western North American dorsally brown coloured frogs, R. pretiosa, R. aurora. Whereas American Wood Frogs hibernate on land as far as is known, the Eurasian brown frogs hibernate often in the water with some species hibernating sometimes on land as well. The five species in the green frog group are primarily pond frogs somewhat like the North American Rana clamitans group in ecology, hibernating most often in the water and with tad- poles that sometimes overwinter. But their patterns often include spotting that resembles North American Rana pipiens, and sometimes a middorsal stipe, a character found predominantly only on the Wood Frog in North America. Included in the Eurasian green frog group is a complex of two species, the Marsh Frog R. ridibunda and Pool Frog, R. lessonae, and a hybrid of the two, the Edible Frog, R. esculenta, a “klepton” which can cross with either to produce both frogs with the genetic consti- tution of the hybrid and also frogs that are genetical- ly identical to one or the other parental species depending on which is involved in a particular cross. Families which do not occur in North America are Discoglossidae (Bombina, 3 species) and Pelodytidae (1 species). Of the salamanders, only one family (Salamandridae, the newts) is shared but none of the three genera represented occur in North America (Salamandra (1 species), Mertensiella (1), and Triturus (7) [although the species in both endemic North American genera, Notophthalmus and Taricha, were at one time included in the latter genus]. The Hynobiidae eco- logically partly replaces the North American endemic families Ambystomatidae and Plethodontidae with species markedly similar in form and colouration to some of these represented by the genera (one species in each) Hynobius, Salamandrella, Ranodon, and Onychodactylus. The book is packed with fascinating comments and rich in data. The historical lore recounted includes that of the ancient aboriginal people of the Kamchatka Peninsula, the Itelmens, who suspected that Siberian Newts, Salamandrella keyserlingii were spies of Gaech, lord of the underground. The Itelmens therefore killed newts in the belief that if 666 the animal saw a man and was allowed to return to its master and report him that man would die (page 8). Another ancient belief recounted is that mermaids by their sad songs decoyed people into deadly swamps, which may have been based on the evening calls of Fire-bellied Toads, Bombina (page 9). The ecological sections of the species accounts include such data as that on the Moor Frog, Rana arvalis, which in one 6 km? area of mixed forest in West Siberia, has a population estimated at approximately 90 000 individuals (biomass 190-200 kg), which will consume 55-60 kg of invertebrates per summer day (page 322). In contrast to North America, where none of our salamanders are known to be freeze- tolerant and all fall far short in ranges of the most northerly ranging frogs, Salamandrella keyserlingii is reported to survive temperatures of -35 to -40 Centigrade, and ranges farther north than any other Russian amphibian. It is the only amphibian to occur THE CANADIAN FIELD-NATURALIST Vol. 116 - in the severe climate of the Kamchatka Peninsula. As in North America, only one frog ranges north- ward over the Arctic circle. In Canada, the species is the American Wood Frog, Rana sylvatica, in Russia the Siberian Wood Frog, Rana amurensus, although the Moor Frog, Rana arvalis, comes close in the eastern portion of the country. If the book has a fault, it is that the text, although having a list of citations at the end of each species account, does not cite individual papers after state- ments in the text, thus not allowing the source of particular data of interest to be quickly traced in a lit- erature that will be so unfamiliar to most North Americans. FRANCIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada The Atlas of the Amphibians and Reptiles of Vermont December, 2001 By James S. Andrews and 1400 dedicated volunteers and organizations. 2002. [Privately printed] James S. Andrews, Middlebury, Vermont. 90 pp. Available from Cindy Brown, Biology Department, Middlebury College, Middlebury, Vermont 05753. U.S. $15 (plus 0.75 Vermont sales tax). On the two occasions in the past year when Jim Andrews has given a presentation at a meeting we have both attended, he invariably has begun by illus- trating the former dependence on non-Vermonters for information on amphibian and reptile distribution in the state by singling out my critical review of a book on New England amphibians and reptiles in 1984 (Canadian Field-Naturalist 98(3): 406-407). To illustrate the lack of a comprehensive check of museum collections by the authors I had appended a note by F. W. Schueler his (until then) unreported and therefore overlooked specimens of Chorus Frogs, Pseudacris triseriata, the first record of the species for New England, collected in 1975 in north- ern Vermont and deposited in the Canadian Museum of Nature. Actually, a distributional database for the state had already begun by the time my comments were pub- lished, thanks to The Nature Conservancy's Heritage Program whose methodology was introduced throughout New England in 1983. An advisory Endangered Species Subcommittee for Reptiles and Amphibians was formed for Vermont with Mark DesMeules as chair. This led to the compilation of a — Preliminary Atlas of the Reptiles and Amphibians of Vermont published in 1995. Included were records from museums, publications, and a host of field observers. The project has continued and since 1995 over 1400 individuals have contributed 12,000 new records, for a database total of over 30,000 records. Jim Andrews (who has contributed over 10,000 records himself) has coordinated the atlas project since 1994 and is the current chair of the Vermont Reptile and Amphibian Scientific Advisory Group. No longer was Vermont largely dependant on occa- sional travellers for information on rarities. The maps presented cover 40 species considered native to the state: 10 salamanders, 11 frogs, 7 tur- tles, 1 lizard, and 11 snakes. For the Blue-spotted Salamander complex multiple maps are included to separate the complex as a whole, each of the two diploid species, and each of the polyploid hybrid combinations associated with each of these. In addi- tion, three salamanders and two turtles are listed for which specimens exist which are thought to be the result of mislabelling or released captives (of these only the Allegheny Mountain Dusky Salamander is mapped). The maps indicate if a species is present in any of approximately 245 geopolitical “town” divi- sions (key to these is given on page 15) by hatching for historic records (prior to 1976) and shading for current records (1976 through 2000). Most impor- tant, and an innovation Canadian atlassing projects might well consider, is a distinction (in this case a dot) added for the most valid records, those which are documented by a photograph, tape recording (for frog calls), or specimen. Inner front and inner and outer back covers present colour-coded maps of numbers of new species reported in each township since the 1995 edition, total species reported for each 2002 township as of 1 January 2001, and degree to which each town has been surveyed to show the areas most in need of additional survey. The simplified mapping system works well for encouraging volunteers to “fill in the blanks” but it is frustrating for any but the broadest-scale zoogeogra- phers and ecologists. Those needing precise plotting of localities for finer scale comparison with vegeta- tion or geographic variables will not find it here. The centre-of-a-block method is arguably less work. It also is conservation-friendly in as much as exact localities are not pin-pointed and thus avoid (or at least make more difficult) its use for the exploitation of rare or commercially valuable species. However, at the same time it obscures critical habitat locations from possible developers (or opposers of develop- ment) who might not destroy populations of rarer species if they could readily check for their presence. Either could, of course, take the additional time nec- essary to access the original data files to check if a rare species indicated for a town might be present in the exact locality of their concern. Hopefully, responsible people would. The concluding text has tables of the number of records for each species and a plea for continuing help as there is so much still to learn. A section on documenting records is the clearest explanation and justification I have seen for why and how to do this with special care. In addition, there are tips on searching and handling, permits and licences Snake By Chris Mattison. 1999. Firefly Books, Willowdale, Ontario. 192 pp., illus. $39.95 This large format (28.7 X 23.7 cm), “coffee table” style book has spectacularly crisp colour photographs, and an encyclopedic text packed with capsule facts. It begins with a single-page introduction which pro- motes an interest in, and appreciation of, snakes. Following are three major divisions. The first, the Essential Snake, covers evolution, environment, size and shape, scales, anatomy and movement, skull and teeth, sense organs, hunting and feeding, venomous snakes, passive defense, active defense, reproduc- tion, conservation, and classification of snakes, most topics covering two facing pages and all illustrated by colour photographs and diagrams. The second, Snake Gallery, covers a selection of 61 primary species each also mostly on a two-page spread, with a center portrait of the featured species sprawled across both pages. Fitted around it are the species name, a summary of its main characteristics, a diagram of relative size giving average and record lengths, additional pictures showing particular mor- BOOK REVIEWS 667 required and ideas for educators. There is an eight- page, four-column list of contributors of up to 19 records, followed by a page listing those contributing 20 to 99 and more than 100. A list of field employ- ees and drift fence volunteers includes 72 people and an Audubon Take PART Program an additional 36; organizations that contributed records totalled 36, and museums 13. There is a five-page bibliography, a list of eight advisors, a page of useful publications, tapes, and websites, three pages of additional reading on amphibians and reptiles, and a two-page reporting form for observations. This and other atlassing projects in neighbouring states and provinces are not just invaluable for the documentation in itself, but also in their promotion of future effort and usefulness as a basis for conser- vation initiatives. Because they specifically acknowl- edge each individual contribution they promote pub- lic support and encourage continued involvement. The present work based on presence/absence data will lead eventually to the next, and even more diffi- cult step in atlassing, the mapping of annual relative abundance geographically. To do this we yet need to learn how to accurately measure amphibian and rep- tile populations, a task both challenging and illusive. FRANCIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa K1P 6P4 Canada phological or behavioral characteristics, a “fact file” giving Latin name, family, habitat, breeding, feed- ing, and distribution, the latter with a tiny globe with the geographic range in red, and an inset box with a topic supplemental to the main text, either conserva- tion issues or other related species. The third, Snake Directory, is an unillustrated four-columns-to-a-page listing. Each family is given with its characteristics followed by included genera in alphabetical order. Each genus has the number of species, size, distribution, habitat, food, reproduc- tion, notes. Species are listed with scientific and English names (if one of the latter is in common use) and a dot if featured in the earlier Snake Gallery sec- tion. The book concludes with a 52-entry Glossary, Index, and Acknowledgments. As a visual of major topics in snake biology and representative world snake diversity along with a convenient easy-to-scan summary of conservative classification and species of world snakes the book ts a success. It will be useful on any bookshelf for quick reference to the diversity of snakes, Unfortunately, it 668 THE CANADIAN FIELD-NATURALIST lacks any bibliography, so the reader can not follow up topics of particular interest to sources where they are discussed in greater detail. For this, reference to more detailed and academic snake texts will be needed. Vol. 116 - FRANIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada A Field Guide to the Amphibians and Reptiles of the Maya World: The Lowlands of Mexico, Northern Guatemala, and Belize By Julian C. Lee. 2000. Comstock Publishing Associates, Cornell University Press, Ithaca, New York. 402 pp., illus. U.S. $35.00 Amphibians and reptiles do not generate the con- stant degree of general field-naturalists' attention, travel, and consequent book demand that birdwatch- ing does and therefore the appearance of a field guide to them for an area south of North America and Europe is an event. A Field Guidé to the Amphibians and Reptiles of the Maya World is such a book. It is of general interest as amphibians and reptiles (in contrast to birds) are non-migratory, except for marine turtles, and therefore only the latter may actually share individuals with the north- ern hemisphere. Other groups have long evolved independently and the southern herpetofaunas have not been periodically eradicated through the Pleistocene by repeated glaciations. Most field guide coverage is defined by continents or “artificially” (from a biological point of view) by political units: countries, states, or provinces. In this guide to the Mayan world, the choice taken breaks from this tradition by selecting a geographical block lying north and northeast of the 600-metre contour in the highlands Alta Verapaz, Guatemala, and the Meseta Central of Chiapas, Mexico. The area includes the southern Mexican states of Yucatan, Quinta Roo, Campeche, and portions of Tabasco and Chiapas, as well as the countries of Belize and Guatemala and portions of Honduras. The lowlands thus focused on were the home of the indigenous people of the Yucatan peninsula, the Maya, hence the book's title. The guide begins with the mandatory Preface and Acknowledgments. These are followed with an Introduction explaining the area covered, how to use this book, conservation, organization of the species accounts. An Environment section outlines physio- geography, climate, and vegetation; a habitats one describes coastal, agricultural, freshwater, forest, and savanna. A discussion of the composition of the her- petofauna precedes the species accounts which are the bulk of the text, 238 pages. The book concludes with a Glossary (11 pages), Literature cited (7) and Index (8). Species-by-species coverage includes paragraphs on identification, similar species, distribution, natu- ral history, and key reference citations for each account. It is richly illustrated. There are 365 figures; 174 of these are black-and-white diagrams (17), habitat photographs (24), drawings of key features; and tadpoles (for frogs and toads) of various species (133) dispersed through the appropriate sections of text. A section inserted between pages 50 and 51 has 181 colour photographs of individual specimens. In addition, there are 163 text maps, one each of geog- raphy and of vegetation, 161 of species distributions with range shown as solid patches. The herpetofauna includes 188 species: 2 caecil- ians, 6 salamanders, 36 frogs and toads, 2 crocodil- ians, 16 turtles (5 marine), 52 lizards, and 74 snakes. The environment is tropical with freezing tempera- tures unknown and a mean annual temperature gener- ally ranging from about 25 to 26°C. Striking in com- parison with herpetofauna in the northern climate of Canada is the presence of the worm-like caecilians and crocodiles, the low numbers of salamanders, and high relative proportion of lizards and snakes. In all of Canada, totals for native species are 21 salaman- ders, 24 frogs and toads, 13 turtles (5 marine), 6 lizards, and 25 snakes (even the addition of species likely introduced adds only one salamander, three turtles, and one lizard). There are few species in com- mon other than four marine turtles, only the Snapping Turtle (Chelydra serpentina), Brown Snake (Storeria dekayi), and Milk Snake (Lampropeltis triangulum), each represented by a different subspecies from Canadian populations. Other genera in common do not share any species: the widespread Bufo, Hyla, Rana, Eumeces, Thamnophis, Nerodia, Coluber, Elaphe, and Crotalus. A major omission is that the contents do not list the species accounts individually nor is any checklist or even keys to the included species provided to sum- marize the species covered. Individual species have to be searched for through the index. The accounts do not reference individual statements to source, and much more informative spot (locality) distribution maps should have been possible for the area. For the south-orientated adventurous tourist-natu- ralist this well-executed and effectively illustrated book provides a welcome guide to amphibians and reptiles and their habitats that may be encountered in 2002 the northern edge of eastern Central America. For the armchair zoogeographer it opens interesting con- trasts to familiar northern species and environment. Either in field jacket or bookshelf it is a reference worth referring to repeatedly. Dinosaurs, The Encyclopedia. Supplement 2 By Donald F. Glut. 2002. McFarland and Company, Jefferson, North Carolina. 685 pp., illus. U.S. $75.00. Donald Glut is a dinosaur fanatic. This is obvious, through his published essays (Jurassic Classics, 2001, McFarland and Company), his growing com- pendiums of dinosaur knowledge (for example, The Dinosaur Dictionary, 1982, The Citadel Press), and the influence on culture these extinct creatures repre- sent (The Dinosaur Scrapbook, 1980, The Citadel Press). In 1997 Glut produced Dinosaurs, The Encyclopedia (1997, McFarland and Company). Its 1076 pages was a considerable undertaking. But the problem with any printed encyclopedia, especially in this age of dot.com, is the speed at which informa- tion is dated. Resolving this, Glut later published a supplement to the encyclopedia (2000, McFarland and Company). And most recently, Dinosaurs, The Encyclopedia, Supplement 2 was issued, foreworded by dinosaur expert Phil Currie. The first hundred pages or so recap and enhance current dinosaur research themes like extinction, the relationships between birds and dinosaurs, and the thermal make-up of these saurians. The bulk of the 687 pages are devoted to descriptions and revisions at the generic level of dinosaurs; new discoveries, new species, and new ideas. And like the first sup- plement, a section at the end gathers the locations of museums and sites where dinosaurs are prominently showcased. This, however, is a subjective list with several sites absent. One must applaud the author’s continued effort to be on top of all that is dinosaurs. Combined, all three The Archaeology of Animal Bones By Terry O’Connor. 2000. Texas A&M University Press, College Station, Texas. 206 pp. $34.95 U.S. Terry O’Connor’s The Archaeology of Animal Bones proves to be a useful guide not only for zooar- chaeologists and even paleontologists alike, but for mainstream archaeologists as well. Non-human remains have the potential to reveal “different forms of data inherent in them” O’Connor maintains, “each BOOK REVIEWS 669 FRANIS R. COOK Researcher Emeritus, Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada titles (all still in print) total some 2200 published pages and cover the popular dinosaurs like Albertosaurus to the less known Zuniceratops. Each generic entry reviews the most recent literature. These are followed by “key references”, thus avoid- ing any misconception of being all encompassing. As one would expect with such a mountainous pro- ject, typographical errors, or simple omissions are to be expected [in a moment of admitted vanity, the first supplement listed Karen Chin erroneously as the sole author of the paper describing the Tyranno- saurus rex coprolite from Saskatchewan]. This aside, included with many of the entries are notes gathered from conversations and lectures, information not usually recorded. In addition, a plethora of pho- tographs and illustrations, many from original publi- cations if not from the author’s own collection accompany the entries. Unlike other substantial and significant compila- tions like The Complete Dinosaur (1997, Indiana University Press), Encyclopedia of Dinosaurs (1997, Academic Press), and The Dinosauria (1990, University of California Press), if Glut is up to the task, his three volumes and hopefully further supple- ments, will be the benchmark of dinosaurian synop- sis. A big proviso, however, would also rest with the publisher — McFarland and Company. They should be recognized for sticking with Glut’s vision and the high quality of the end product. TIM TOKARYK Box 163, Eastend, Saskatchewan SON OTO Canada of which conveys information about the original ani- mal, its life and death, its interface with people, and the complex events and processes which have led to its survival as an archaeological specimen” (page 53). The author covers the breadth of themes in sequence; from taphonomy, excavation, identification, and quan- tification of specimens, to age at the time of death, pathology, environmental influences and signatures, 670 to human interaction. The text is straightforward and easy to read, heavily illustrated, and referenced, useful for an introduction to the field. The Archaeology of Animal Bones confesses to be as broad based as possible in the many approaches to the study of associated bones in archaeological sites, yet it is produced through the experiences of its author. In interpreting body-part analysis for exam- ple, O’Connor notes “there is no ‘right way’ to do it, but by applying a wide range of procedures to the same data sets, we can at least see which patterns in the data are inherent to those data, and which are characteristics of a particular analytical technique” (page 79). This is true for much of the methodology in acquiring the data and their interpretation. BOTANY THE CANADIAN FIELD-NATURALIST ——— a TNT ET CO Vol. 116 - Inadvertently, The Archaeology of Animal Bones would be a suitable companion to Steve Roskams British oriented Excavation (Cambridge University Press, 2001), though the latter is part of a larger series of archaeological manuals. Admittedly, O’Connor’s volume is rich in non-North American examples with modest references to North American sites and research. Yet the clear, step-wise approach by the author, and the broad scientific appeal of the volume makes it highly effective in illustrating the potential of vertebrates in telling a story of the past. TIM TOKARYK Box 163, Eastend, Saskatchewan SON OTO Canada Rare Native Vascular Plants of British Columbia, Second Edition By George W. Douglas, Del Meidinger and Jenifer L. Penny. 2002. Crown Publications, 521 Fort St., Victoria, British Columbia V8W 1E7, Canada. 358 pp. $45.00 + GST and postage. The previous edition which was published in 1998 contained information with distribution maps and excellent line drawings, habitat/range information, Global/Provincial Rank, and status Red or Blue. The Red List included 193 taxa that were candidates for legal designation as endangered or threatened and 42 historic or extirpated taxa plus seven not considered as above but included in the Red List. The Blue List included 357 taxa that could become candidates for the Red List. In this new edition considerably more information is provided in the Habitat/Range section. The Red List has been increased to 271 taxa and the Blue List to Flore printaniére By Gisele Lamoureux with Roger Larose photograph col- laborator. 2002. Fleurbec, 198 chemin Grande-Grillade, Saint-Henri-de-Lévis, Québec, GOR 3E0 Canada. 576 pp. $29.95, including GST and postage. This soft cover book on the spring flowers of the province of Quebec has an absolutely beautiful col- lection of colour photographs which were taken by 15 photographers: Gisele Lamoureux (187), Roger Larose (59), Francis Boudreau (12), Camille Rousseau (7), Michel Boulianne (3), Jacques Labrecque (2), Normand Lalonde (2), Claude Allard (1), Yves Boudreault (1), Line Couillard (1), Lucette 346 taxa. In addition, many of the maps have been updated and some sites that were previously marked by a black dot have been changed to a black circle to indicate that a taxon has been extirpated or not known from that area for a long time. Additional notes have frequently been added for many of the taxa. This publication is a tremendous step ahead for the knowledge of the vascular plant flora of the province of British Columbia. I would be more than happy to see similar updated works for any of the other Canadian provinces. WILLIAM J. CODY Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada. Durand (1), Alain Giard (1), Miroslav Grandtner (1), and Sylvain Lamoureux (1). The text of this volume is written in French. The only exceptions to this are an English common name for each species and of course, the scientific names which are in Latin. The first page describes the joy of seeing the beautiful wild flowers which appear in the spring. This is followed by a list of the common names and page numbers as they appear in the text, introductory information including North American floristic habitats and vegetation zones in Quebec and Labrador. 2002 The first section provides two pages for each of 105 wild flowers. The left page is initiated by the French common name in large bold face, followed by an English common name in small face, addition- al French names in small face, and the scientific name and synonymy with information on the scien- tific names. This is followed by a description of the plant, its habitats, and other information all of which is wrapped around a small distribution map showing where it occurs in North America, and facing its beautiful picture on the right hand page. Then comes a second section which includes eleven pages depict- ing flowering trees, and a third section, in the same Ecological Management of Agricultural Weeds By M. Liebman, C.L. Mohler, and C.P. Staver. 2001. Cambridge University Press, New York. xi + 532 pp., illus. U.S. $120. Agriculture, as cropland and/or pasture, is found in some form whereever mankind settles. A major challenge for agricultural production has been the existence of unwanted plants (weeds) within the stand of desirable plants (the crop). Within the developed world herbicides have been the major method of control and key component of textbooks on weed management. As a result of increased inter- est in invasive plant species, societal demands for a cleaner environment and sustainable agriculture, non-chemical methods of control are gaining societal support. An indication of such support can be found in the suggestion from the Johannesburg World Summit 2002 indicating dependence on chemicals must be decreased and a more ecological approach to food production be developed. Liebman et al. have written this book specifically focusing on eco- logically based methods of management. In the words of the authors they were “able to provide in- depth treatment of subjects that most weed science books treat only briefly.” This in-depth treatment of subjects is done in 11 chapters organized into three main sections. The three main sections focus on: the need for and approach to ecological management; impact of the environment and disturbance; weed evolution, and plant community structure. Each of the chapters has references lists, with some exceeding 250. A BOOK REVIEWS 671 order as Section One with continuing information on the species and many additional photographs. The volume is completed with a glossary, a most useful bibliography, an index to other Fleurbec guides, and an index to this volume. Overall, this is a most beautiful volume. WILLIAM J. Copy Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada sampling of what the chapters include follows: mechanical weed control, increasing crop diversity, biocontrol with livestock grazing, insects and pathogens, and modification of cropping systems to enhance the crops’ competitive ability. The contents rely on basic ecology, plant biology, and applied ecology literature. The authors put forward three objectives for eco- logical management of weeds: (1) weed densities should be reduced to tolerable levels not eliminated; (2) amount of damage to the associated crop should be reduced with a given weed density; (3) shifting of the weed species or community to a less aggressive and less persistent composition. Having provided the objectives, the authors also supply areas of knowl- edge needing further research to help meet the objectives. Ecological Management of Agricultural Weeds in this reviewer's opinion should make an ideal text for an upper year undergraduate or graduate level course. The well researched contents with extensive references could easily provide a good grounding in the subject matter. With increased societal demands to reduce reliance on herbicides and for greater reliance on ecological approaches, the demand for this textbook can only increase. I congratulate the authors in a well timed and well done text. M. P. SCHELLENBERG 434 4" Avenue SE, Swift Current, Saskatchewan S9H 3M1 Canada 672 Fungal Conservation: Issues and Solutions Edited by David Moore, Marijke M. Nauta, Shelley E. Evans, and Maurice Rotheroe. 2001. Cambridge University Press, Cambridge, UK and New York. x + 262 pp., illus. U.S. $95.00. The realization that fungal species (with few exceptions the book concentrates on the macrofungi, such as mushrooms and allies) are experiencing sig- nificant declines has made conservation a major con- cern. Fungi are ubiquitous, they make up a signifi- cant part of the microflora in soils, and the roots of most higher plants have a beneficial association (mycorrhizae) with the fungi. Key words and phrases from some of the chapter titles are: trends and perspectives, nature manage- ment, forest fungi, ecosystem, future of fungi, fungi as indicators, threats to biodiversity, valuable, rural economies, survey, national mycological societies, and optimism. Economically the wild mushroom industry is mushrooming. In 1992, nearly 2 million kg were har- vested in the Pacific Northwest of the USA, bringing over US$40 million to the economy (Molina et al.). And the global trade in matsutake alone is estimated at US$3 to 5 billion annually; for chanterelles it is about US$1.5 billion (Aurora). In some communities in China, Bulgaria, Italy, and Zimbabwe wild mush- rooms provide 50 to 100% of a family’s annual income (Aurora). In Champa (Tibet) the intact, healthy forests are perceived as the key to rural development rather than as an impediment to it (Aurora). Thus sustainable management of the wild mushrooms is critical to the welfare of many towns and villages. Regarding sustainability of harvests, Aurora comments that wild mushrooms, like pine nuts and huckleberries, can be harvested without sig- nificantly damaging our forests and that buying them in a gourmet restaurant probably comes at less cost to biodiversity than wine, beef, or almost any other item on the menu. From committees focusing on global conserva- tion to the effects of wild mushrooms on rural economies, the 22 chapters in this book provide an introduction to the varied problems and proposed solutions to fungal conservation. In the introductory chapter the editors present an overview of the threat to microorganisms and establish that fungi are important. In giving readers perspective on their importance, Staley (1997, Current Opinion in Biotechnology 8: 340-345) is quoted “the tree of life is largely a tree of microorganisms ... much of the diversity on Earth is microbial with the plants and animals appearing as small, terminal branches.” Before solutions can be implemented it is essential THE CANADIAN FIELD-NATURALIST Vol. 116 to define the problem, and in the case of fungi this has not been easy. Several authors emphasize that effective conservation programs need the input of mycologists (those who know and study fungi) and mycologists are seen in many parts of the world as an endangered species. Public education is discussed as the key to gaining the necessary resources and this is a long term effort. In the more immediate future, Molina et al. argue for the development of a cadre of parataxonomists (those of less formal mycology schooling but trained in fungal identification and methods of study) that can evaluate the mycological- ly valuable sites. Courtecuisse, in a chapter titled “Current trends and perspectives for the global conservation, of fungi” gives an overview of the main tools being used by fungal conservationists; i.e., inventories, mapping programs, and Red Data Lists. And these items appear in various forms in all chapters. Several authors bemoan the lack of base-line data on which to judge the present status of diversity or frequency. Jalink and Nauta show how such data can be used to calculate the mycological value of a site. At least 20 European countries have Red Lists for fungi (essen- tially mushrooms and allies) and, although the crite- ria for listing may differ, these have been valuable in managing species (Arnolds). Arnolds in an intrigu- ing chapter titled “The Future of Fungi in Europe” presents a wide ranging discussion of many facets of fungal conservation. His discussion of checklists concludes with “Any checklist is better than no checklist.” Nota bene colleagues who hesitate! It is disappointing that only one contribution focuses on the Western Hemisphere. However, that seems to reflect the importance given to fungal con- servation in the area. Molina et al., in a detailed, thoughtful presentation, deal with the problems, logistical and legal, of conservation in the United States’ Pacific Northwest. The presentation con- cludes with eight guidelines for mycologists to con- sider when embarking on fungal conservation efforts. This is a well written book, broad in mycological scope, and the topics are concisely presented. It is a good starting point for anyone interested in fungal conservation and should be required reading for North American ecologists, plant diversity researchers, land managers, and foresters. J. GINNS 1970 Sutherland Road, Penticton, British Columbia V2A 8T8 Canada 2002 ENVIRONMENT BOOK REVIEWS 673 Two Hundred Years of Ecosystem and Food Web Changes in the Quoddy Region, Outer Bay of Fundy By H. Lotze and I. Milewski. 2001. Conservation Council of New Brunswick, Fredericton. 188 pp. $30. The Bay of Fundy is a Canadian and international jewel. This book presents one of the best and most updated compilations and analyses of various rele- vant topics for a fascinating, outstanding, and very productive ecosystem, the Quoddy Region. Besides classical subjects such as fish, sea mammals, birds, and oceanography, this landmark publication even adds several new topics of interest for the ecological integrity of “the Bay”, such as archeology (humans), benthos, marine plants, plankton, species extinction, whaling, and sealing. Many indicators of environmental health and ecosystem integrity are thoroughly investigated by the authors. All indicators convincingly show that the Canadian North Atlantic was treated as badly as any other of the major fishing grounds in the world; the Bay of Fundy and its Quoddy Region are no exception. The authors present compelling, if not final, evidence for this fact. Lobster catches steeply increased in Eastern Canada during the last years, providing the American market with a highly-priced commodity; it is very hard to believe that this har- vest could be on a sustainable level. This book is another unique compilation of inter- preted information and data sources proving con- vincingly how human activities have altered the planet. From the Passenger Pigeon, over the Labrador Duck and Great Auk to the Sea Mink; all of these animals were creatures of the Bay of Fundy region that were wiped out by poor management of natural resources and by human greed within few generations. However, when taking into considera- tion the Canadian realities such situations should perhaps not be judged too easily. Harsh environ- ments deserve different perceptions and judgments, different than those that are easily done in conve- nient cities and offices. This appears to be a slight weakness of this book: the authors see changes in the light of good or bad, right or wrong. Most population declines are “bad” and increases are “good”, or they are not further discussed. The overall paradigm of the book that “native species are good, Europeans are evil”, could be put into doubt. In contrast to what is presented in the book, it could be possible that ani- mals in an “undisturbed” environment (already a debatable term as such) were only abundant in low numbers, and that population increases, often inter- preted as a “recovery” or safe situation, present something very bad. The current Atlantic-wide extension of the Northern Gannet population is such an example, increasing Cormorant numbers in the Bay of Fundy are another one. An increase is not always good or natural; it is well known that marine habitat degradation such as overfishing can trigger the high abundance of small prey, which increases many waterbird populations. Small-scale signals, such as found in the Bay of Fundy Quoddy Region, need to be seen and interpreted in their global con- text. The question why Atlantic Right Whale popula- tions, fully protected for many years now, are still not “exploding” remains unanswered. The authors provide a powerful message on how the Bay of Fundy region was altered in the last two hundred years. The decline of herring in the Bay of Fundy is considered already a traditional trend. Here another supporting argument to underline these facts: harvesting down the food chain has now reached the plankton and rockweed level. The promotion of pure economic growth is of no value to anybody who needs to make a sustainable living from the Bay of Fundy and its resources. Besides compiling many references and informa- tion about food webs and the ecosystem of the Quoddy Bay, the authors also present very interest- ing reviews on whaling and sealing. It would be great to make the wonderful literature reference selection (24 pages strong), and the compiled raw data, accessible to anybody; e.g., via a database on the Internet/(WWW. This book demonstrates the tremendous value of long-term data sets. Unfortunately, for most impor- tant management questions such as populations of fish and bird species or habitat quality monitoring, these databases are missing, were not derived from a rigorous statistical design, or were never maintained for the entire Bay of Fundy. However, such high quality monitoring data are still needed and a crucial requirement for any sound management, and certain- ly for a sustainable nation. Instead, we are forced in many cases to make interpretations from commercial fisheries catching and harvest data. These ones are really flawed (e.g., by technology and market demand) and therefore they are almost not inter- pretable, presenting nothing more than pseudonum- bers of low ecological value. The book compensates for this problem somehow by analysing as many data sets as possible. The book shows that success stories regarding the poor state of the Bay of Fundy’s health are few, and normally driven by the need of managers to produce “progress”. For instance, stocking fish is not a solu- tion at all; this is well proven elsewhere and even considered to be harmful for the fish stocks as well as for the habitats. Other “success stories” have reached over the years a point that they become ironic; e.g., recovery of groundfish, international 674 conservation issues of the St. Croix river, and dams affecting runs of Gaspereau and salmon near Moncton or in the St. John river. If the dams were gone (or opened), the fish would do fine. The contributions of this book are manifold: small formatting and figure errors can easily be forgiven, even that a U.S. perspective is not included (the Quoddy Region has a substantial U.S. border). After the publication of this book, it becomes obvious that the science is done, what’s needed now is action. Boldly spoken, the Irving company is still running its refinery and pulp and paper mill in the traditional way, which contributes to the reputation of St. John’s as the “armpit” of Canada. But no wonder when an industry is allowed to monopolize the regional news- papers and the political and job scene. To the general surprise of the many (international) visitors, pulp and paper mills are still releasing their waste water in the rivers and Bay of Fundy, a nuclear power plant is located right at the waterfront, and aquaculture is pro- moted. Although the city of Moncton had severe E. coli outbreaks, there is still no major sewage treatment Autumn, A Season of Change By Peter J. Marchard. 2000. University Press of New England, Hanover, New Hampshire. 151 pp., illus. + plates. Cloth U.S.$50; paper U.S.$17.95. This book, by field biologist and biology professor Peter J. Marchand, is a remarkable natural history work. It takes one season, autumn, and looks at it from such diverse and fascinating angles — some which never would have occurred to me — that my head is still reeling from all I’ve learned. That’s not surprising. I couldn’t have known about all those things because I’m not a scientist. But Marchand is, and he’s an awfully good writer too (and philosopher, and photographer), who manages to render complex chemical and biological processes understandable to a non-scientist like me. In the book, Marchand focusses on the terrestrial environments of his personal experience. He illumi- nates various autumn phenomena through fifteen essays on topics such as freezing, senescence, dispersal, migration, the fall rut, autumn in ponds, harvesting activities, and more. I discovered all sorts of fascinating things in this book. From the wide array of light-sensitive pig- ments in plants, to the biochemistry behind fall colours. From the diversity of molting processes in mammals and birds, to leaf self-amputation. From intracellular ice formation in plants, to freeze avoid- ance and tolerance in insects. From the factors prompting bird migration, to the impressive journeys of insects. From the stakes involved in the fall rut, to THE CANADIAN FIELD-NATURALIST oe mn oe a TG. = = TC Vol. 116 - approach put forward for the watershed of the Bay of Fundy. This should be embarrassing to any Canadian citizen, and certainly to any informed conservationist, manager and politician, to say the least. It is fortunate that this publication was made possible by different (private) funding agencies and NGOs (e.g., Conservation Council of New Brunswick). The authors can be congratulated for their achievements. They conclude that “there is no evidence of an ecolog- ical simplification of the entire food web defined as the loss of an entire trophic level or functional group. But, the food web is severely altered and we find strong shifts in species composition and dominance patterns that affect predator-prey, habitat and competi- tive relationships”. It is now up to us to restore and to save the Bay of Fundy from even more harm. FALK HUETTMANN Postdoctoral Killam Fellow, Geography Department, Uni- versity of Calgary, Calgary, Alberta T2N 1N4 Canada. E- mail: falk@ ucalgary.ca the risks of hibernation. From amazing fall harvest accomplishments among mammals, to the highly involved process of leaf decomposition. Marchand the poet and philosopher comes out in passages like “For the apparent disappearance of many plants and animals, autumn is often seen as an end. But the seasons are part of a continuum, a revolving process of birth, death and renewal — and if such could be said to have any beginning or end, then fall could just as well be viewed as a beginning.” Speaking of poetry and philosophy, Marchand sprinkles his essays with passages from the works of late great nature writers such as John Muir, Henry David Thoreau, and John Burroughs. And the author’s own black-and-white photographs add addi- tional touches of creativity. As Marchand writes in the preface, “while the book is primarily concerned with nature and science, it also about art, literature, and a smattering of philosophy, because to discuss science in isolation from these would be something like studying the life of a flying squirrel in a cage; fruitful to a point, but missing essential context.” Marchand creates a most fascinating autumn con- text in this book — one that both delights and teaches. Autumn, A Season of Change is certain to become a well-thumbed resource in my natural history library. R. SANDER-REGIER R.R.5 Shawville, Quebec JOX 2Y0 Canada 2002 BOOK REVIEWS 675 Making Parks Work: Strategies for Preserving Tropical Nature Edited by J. Terborgh, C. van Schaik, L. Davenport, and M. Rao. 2002. Island Press, Washington. xix + 511 pp., illus. Cloth U.S. $65; paper U.S.$32.50. This book contains 32 chapters by a variety of authors on management of parks in the tropics. These result from a meeting in Florida in August 1999 of 30 tropical park conservation experts. The book has a purpose as a reference guide for field workers. The book is divided into four parts: (i) introduction — 3 chapters; (ii) case histories — 14 chapters; (iii) themes — 13 chapters; and (iv) conclu- sions — 2 chapters. The introduction defines the need for parks. The overall theme is that extinction rates are growing continuously under increasing population growth, especially in tropical regions. Parks are essential for the protection of many of the more susceptible species. The most ecologically diverse areas are always the most fertile areas, which are thus most in demand for agriculture. The most critical species are often the larger animals, which usually require larger areas to graze or to hunt and are usually less tolerant of human interference. Very strong opinions are expressed that attempts to integrate conservation and sustainable development just do not work. The con- cept of the “noble savage” with an innate desire to conserve nature, is also debunked. These are both very pertinent to me, as I am reviewing this book while attending a workshop, sponsored by the World Bank, relating to management of endangered envi- ronments in tropical Africa. The workshop is taking place in Yankari National Park, Nigeria, where I can listen to elephants and lions, as well as other large animals, in the evening. A large part of the discus- sion relates to involving local villagers in sustainable development and environmental management. The concepts that this book is opposed to are being strongly endorsed at this workshop. I am not sure who is right, but it does seem the sustainable devel- opment concept is crumbling under Nigeria’s rapidly expanding population with the highest priority to many being to keep from starving. I do agree with the conclusion that attitudes (at all levels of society) and institutional capacity are critical to the success of tropical parks, but am less in agreement with the concept put forward that foreign investment, with financial compensation for loss of land, is a good way to resolve the problem. Many case histories are presented from around the tropics. I found these very informative with useful examples of what has succeeded and what has failed. It is also interesting how very different methods suc- ceed or fail in different societies. For me, it was dis- appointing to not see any examples from Nigeria — the country with the largest human population and most diverse environment, as well as some of the most environmental loss in all of Africa. The only West African examples were all related to the rain forest environment and primate species. Yankari NP, where I was sitting to read this, is savannah, but still has many recent local extirpations, some major habi- tats and wildlife species, as well as significant human-wildlife interactions. The elephants have found a liking for watermelons. I have not seen one eat a watermelon yet, but can picture some comical approaches. They have also discovered that they are bigger than the farmers who try to chase them off. This can have disastrous results. The case histories include five from Africa, four from Latin America, and four from Asia. Interesting are the case histories where parks have succeeded. The most intriguing are the private parks of Costa Rica, deriving success largely from ecotourism values. The third section of the book, on Themes, has some excellent examples and recommendations. Very worthwhile topics for tropical park planners include mitigating human-wildlife conflicts, enforce- ment (which is generally felt to not work), eco- tourism, financing, and monitoring conservation efforts. For my work in protecting African parks and envi- ronment, this is a most useful reference. It might be interesting to those involved in park conservation elsewhere or just interested in the tropical environ- ment, but it is rather specialized and probably of lim- ited interest to the average Canadian field-naturalist. WILSON EEDY Geomatics Nigeria Limited, c/o Terfa Inc., R.R.# 1, Glen- cairn, Ontario LOM 1 KO Canada 676 THE CANADIAN FIELD-NATURALIST Vol. 116 - Thoreau’s Country: Journey Through a Transformed Landscape By David R. Foster. 2001. Harvard University Press, Cambridge, Massachusetts. 270 pp, illus. U.S.$16. Naturalist and author Henry David Thoreau (1817— 1862) is revered today as one of America’s first envi- ronmentalists. His journals of his days in the fields and backwoods of early nineteenth century New England were shaped and informed by an ecological consciousness that was unusual for the time and place. Indeed, his observations — and more importantly, his musings — have inspired all kinds of conservation activities in his name. But as David Foster, Director of the Harvard Forest, argues in Thoreau’s Country, the world that Thoreau painstakingly documented in Walden and other works is no longer recognizable. The pastoral setting is now a forested landscape — quite different from the countryside that Thoreau once understood and appreciated. Foster tells the story of this transformation through a generous mixture of quotations from Thoreau’s journals and a series of reflective, engag- ing essays on a variety of topics (meadows, wood lots, fires, animals, and even fences). In reading the sections, it quickly becomes apparent that Thoreau was actually describing a landscape with a heavy human footprint. It is also clear that the New England woodlands today are closer to the kind of landscape commonly identified with Thoreau, but ironically, not the one he intimately knew and enjoyed. As Thoreau himself once observed, “man’s works fall into nature.” BILL WAISER Department of History, University of Saskatchewan, Saska- toon, Saskatchewan S7N 5A5 Canada Voices for the Watershed: Environmenta! Issues in the Great Lakes — St. Lawrence Drainage Basin Edited by G. G. Beck and B. Littlejohn. 2000. McGill- Queen's University Press, Montreal and Kingston, For the Wildlands League, Canadian Parks and Wilderness Society, Toronto. 299 pp., illus. $39.95. Having been involved in a task on the biodiversity of the fishes of the Great Lakes, I assume it was the subtitle of this book that caught my attention. My overly narrow view led me to be unprepared for the nature of the book. The book is an outgrowth of par- ticipation by Littlejohn, ostensibly as a photogra- pher, on a cruise of a U.S. research vessel working on the St. Lawrence River and the Great Lakes. The purpose of the cruise was both study and education, and it involved a number of mainly earth scientists with a breadth of professional interests. The concept for the book involved the following points: (1) scien- tific knowledge made available to the public in an understandable style; (2) a complete watershed approach: (3) a cross-boundary effort involving writ- ers from various U.S. states, Ontario, and Quebec, with an awareness of the impact on the watershed of what is done, and is not done; (4) wide range of top- ics, the bad news and the good; (5) public awareness of, and identification with, the watershed and specif- ic parts of it; (6) scientific, ecological, conservation, and restoration activities; (7) broadscale citizen involvement to improve the health of the watershed. It is difficult to review the book other than in a gen- eral way. It includes a brief Foreword by Monty Hummel of World Wildlife Fund Canada. The bulk of the book is organized in two Sections. Section 1, Voices for the Watershed, begins with a brief Prologue by Bruce Conn emphasizing science and society working together for environmental under- standing. This is followed by a more extensive intro- duction by the editors. The articles in Section 1 are organized under headings Headwaters and Uplands Regions, The Great Lakes, The St. Lawrence River, and Watershed Perspectives. This section includes 31 articles, divided by both subject and geography. The length of individual articles varies from 2 to 2'/) pages (Rescuing Native Fish and The Acid Rain Story) to an average of 5 to 8 pages. Section 2 is headed Finding a Voice. That section includes an article entitled Revelations: The Evolution of an Environmental Ethic and Career, by Michael Keating (33 pages). An Epilogue by Bruce Littlejohn (11 pages), and the Appendices are included in this Section. The latter involve a list of environmental agencies, references, brief author biographies, acknowledgments, and index. There is a high quality and very detailed map of the watershed at the beginning of the book. It, and some of the articles, emphasize the more recent recog- nition of the reality of the Finger Lakes, Adirondack Park, and Lake Champlain system as parts of the 7 Great Lakes-St. Lawrence Watershed. The editors took pains to emphasize that the intent of the project was to reach far beyond the level of providing information, and expressed the hope that the numerous “voices” will motivate and inspire the 2002 reader, as an individual or as part of a group, to become actively involved in helping to protect the watershed and drainage system. They stated that the book is not intended for environmental experts or scientists. They suggested that for that reason the authors are largely those trained to write for the gen- eral public. I was very pleased to see the emphasis by John Jackson on “Bi-National Citizen Action”. Working with the Natural Resource agencies of the various political areas involved in the watershed too often reveals a tendency for each to go its own way rather than recognize they are all involved in the concern for, and the stewardship of, a single, shared ecosystem. The Editors and authors of this book have achieved, to a moderate extent, their stated goals since they rightly insist the future is up to the response of the citizens. I enjoyed the suggestion of Lavigne and Gates that all of us need to know our “Ecological Address” which is somewhat akin to our Conservation of Exploited Species Edited by John D. Reynolds, Georgina M. Mace, Kent H. Redford, and John G. Robinson. 2001. Conservation Biology. Number 6. Cambridge University Press, New York; Wildlife Conservation Society; and The Zoological Society of London. xx + 524 pp., illus. Cloth U.S. $120;, paper U.S.$44.95. What happens when we try to redress the harm we have already done to an exploited species by initiating conservation measures? The use of wildlife for food and other human needs poses one of the greatest threats to the conservation of biodi- versity. However, wildlife exploitation is also criti- cally important for subsistence and commerce by many people. In this text book international experts examine interactions between the biology of wildlife and the divergent goals of people involved in hunting, fishing, gathering, and culling wildlife. Many theories are presented showing how sustain- able exploitation is tied to the study of population dynamics, with direct links to reproductive rates, life histories, behaviour; and ecology. Unfortu- nately many of the theories are rarely put into prac- tice to achieve sustainable use and successful con- servation. For instance, in the Arctic it is difficult to monitor animal population decline and set legal limits for hunting, which are usually determined when a population has already declined. There is growing use of hunters’ knowledge though, and at least they are an effective voice in Arctic wildlife conservation. There is a dramatic change in a whole community BOOK REVIEWS 677 Individual Environmental Footprint. There are many citizens and students only partly aware of the eco- logical problems of our waters and of his/her respon- sibilities in that regard. The size of this book may daunt some potential readers. Others may be discouraged by the unusual nature and brevity of a number of the articles. However, if they read selectively, emphasizing the Introduction and Prologue, they will understand much better their responsibility and role in the protection and wise use of this extremely important freshwater resource. In that case the Editors and authors will have reached the goals we share with them. E. J. CROSSMAN Curator Emeritus (Ichthyology), Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, and Professor Emeritus (Zoology), University of Toronto, Toronto, Ontario MSS 2C6 Canada when a species becomes extinct or endangered. It changes the dynamics of population growth for the endangered species, and there is a cascading effect. One example is, for instance, sea otters which eat sea urchins which in turn eat fleshy algae. Lose the otters and the increased population of sea urchins will kill the kelp forest. Again, problems occur when there is increased agriculture on land outside a wild game reserve. Since this land is usually marginally productive, the land further degrades. A more subtle cascading effect occurs if the population of bees is reduced. Only the most nectar laden flowers get pol- linated and “low-ranked flowers” will not be polli- nated. If wildlife hunting is controlled, hunters will select the biggest and best specimen to shoot. Over time this results in the species becoming smaller through the loss of genes and body mass. An inter- esting chapter gives the results of a study of sustain- able use and control of kangaroos in Australia. Kangaroos have been aggressively hunted on sheep ranches, but the study shows that it is more prof- itable to farm kangaroos and sell their meat and hides than to raise sheep! Even though this book is meant for biology stu- dents, it has much to interest anyone working in wildlife conservation. The moral seems to be “Conservator Beware”! JANE E. ATKINSON 255 Malcolm Circle, Dorval, Quebec H9S 1T6 Canada 678 THE CANADIAN FIELD-NATURALIST Vol. 116 Politics of the Wild: Canada and Endangered Species Karen Beazley and Robert Boardman. 2001. Oxford Uni- versity Press, Toronto. vii + 254 pp. $27.95. As the title suggests this book focuses on the poli- tics of endangered species conservation in Canada although it does not neglect the scientific back- ground of threats and conservation needs of species at risk. It was written under the shadow of the feder- al government’s second failed attempt to deliver endangered species legislation and is therefore in some ways already outdated. Nonetheless it provides a fascinating analysis of the social and political com- plexities of endangered species conservation which are no less challenging now than they were before the third attempt was passed by the House of Com- mons in June 2002. (It subsequently received Royal Assent December 2002 and will come into force by an order in council in 2003) The book is divided into two parts entitled “From Philosophy to Science” and “From Science to Policy”. In the first chapter of part one, co-editor Beazley provides a clear summary of the various philosophical arguments in favour of protecting endangered species, and explains the strengths and weaknesses of each argument. This is followed by a useful but uninspired summary of the, then current, status of species at risk in Canada. The third chapter, on the importance of habitats and ecosystems, is the most disappointing, being almost incoherent in places. An appendix listing the legislation, policy and programs relevant to endangered species for the federal and each territorial and provincial govern- ment is somewhat useful though apparently incom- plete. Policies and programs are only listed for British Columbia and Alberta. An additional appendix apparently summarising a gap analysis report provides neither a reference to the original report nor definitions of key jargon. The last two chapters in part one review the history and current status of terrestrial and marine protected areas respectively. Part two forges bravely into the murky waters of politics. The first two chapters in this section exam- MISCELLANEOUS On Her Own Terms: ine the history of endangered species conservation and politics in Canada, and the federal government’s efforts to create federal endangered species legisla- tion. Of particular interest in the former is the histori- cal trend towards increasing application of science to species designation, recovery efforts, and systematic habitat protection. The latter chapter explains clearly the role of various players in negotiating legislation including an assessment of the effect of scientific opinion (minimal). It also provides an intriguing table comparing the two defunct bills (C-65 and C-33) on the basis of their characteristics and how NGO’s (Non-Governmental Organizations) landowners and industry, and the provinces viewed these characteristics. It would, of course, be nice to have the current bill included in this comparison but that will have to wait for a future publication. The next two chapters take an international perspective. In the first, the politics of species at risk is compared among OECD (Organization for Economic Co-operation and Development) countries with Canada being most similar to Australia but all countries sharing many of the same difficulties. The subsequent chapter addresses the intriguing question of how signing international agreements affects domestic policy with CITES and the Convention on Biological Diversity used as examples. There is no simple answer here. The final chapter provides a summary overview of the complex political process- es discussed throughout the book and concludes that these processes will continue to shape endangered species conservation in the future. I recommend this book to anyone working active- ly in wildlife conservation. It covers issues often thought of as peripheral to the biology of conserva- tion yet critical to the success of any recovery program. CAROLYN SEBURN Seburn Ecological Services, 920 Mussell Road, R.R. 1, Oxford Mills, Ontario KOG 1S0 Canada Annie Montague Alexander and the Rise of Science in the American West By Barbara R. Stein. 2001. University of California Press, Berkeley, 380 pp., illus. U.S. $35.00. Early 20" century was a time of major develop- ment in terms of technology, education, and human settlement. These were especially prevalent in the North American West Coast. With the establishment of new communities in California and the prosperity therein gained, culture and education also emerged alongside. One example of the philanthropic nature supporting the cultural and educational enhancement is reflected in the life long contributions of Annie Alexander (1867-1950), daughter of the Hawaiian 2002 sugar cane baron Samuel Alexander. Her contribu- tions, however, exceeded most other benevolent mag- nates in that her support for research activities at the University of California also included a more practical nature including the personal collecting, identification of thousands of floral and faunal specimens, extinct and extant. Barbara Stein’s On Her Own Terms is more of Alexander’s accomplishments as a contributor to sci- ence and education rather than an all-encompassing biographical tome. For that of a personal nature (for example, her relationship to her companion and fel- low collector Louise Kellogg), one is left with too many assumptions as all surviving documents are of a professional composition. Yet Stein, maintaining her focus, is able to document the vastness of Alexander’s knowledge, experiences, as a woman scientist who “lived passionately, and perhaps most importantly . . . [dealing] with the world on her own terms” (page xiv). Being the daughter of wealth allowed her the opportunity and means to enhance and experience natural history from a practical, first hand view. One of her early adventures while in her 30s allowed her to partake in an African hunting safari with her somewhat unconventional father. Her father’s view of Annie was of an intellectual equal on these ven- tures; a participant rather than an escorted tourist. Resources also allowed her to attend university classes as a “visitor”, thus enhancing her own inter- ests. The untimely death of Annie’s father on the African safari, however, left a void that was only filled by following through on a vision; first, the establishment of a natural history museum on the campus of the University of California. Its mandate would be for research and scientific publication. The nucleus of its collection would be her 3400 speci- mens and if the university would pay for the build- ing, she would pay for salaries and expeditions. This was keenly played on Annie’s part as it allowed her to have control over the research. The 3400 speci- mens and the thousands more she would contribute directly or through funding expeditions, has made the Museum of Vertebrate Zoology one of the most impressive collections in North America. Her interest in contemporary fauna and flora yield- ed only to her interest in fossils. Her participation in BOOK REVIEWS 679 the collection of Mesozoic vertebrates, for example, left her hoping that “I wish my work went further than to simply get the fossils to the university. I should like to follow the saurian . . . to the bitter end, chisel him out of the rock and write learned treatises on his venerable anatomy” (page 19). Her fondness for paleontology would have blossomed further, even after founding the University of California Museum of Paleontoiogy, if it were not for the personal embat- tlements between her and the administration. The university’s initial lack of interest would lead her to exclaim, “paleontology is a foundation science yet it takes a rather spectacular quest . . . to arouse the interest of the [university] administration” (page 180). Still, she contributed specimens, and salaries and financed expeditions. A beckoning question is why a broader audience does not know her? How many individuals in the last century contributed thousands of dollars to one research facility, and thousands of personally col- lected specimens spanning the breadth of natural history? The bulk of Annie’s anonymity comes from Annie Alexander herself. All her donations were to be kept as confidential as possible. As Stein surmises “Alexander was raised in an atmosphere where benevolence was expected but not touted” (page 117). It is unlikely that her anonymity was in any way linked to her own intellectual insecurities (though all admitted that she was of highest ranking among field biologists). “While she felt passionate about science”, Stein writes, “she clearly seemed to favor physical activities over those that were cere- bral” (page 92). On Her Own Terms is definitely worth reading. A pioneering, early 20" century woman making waves in the male-dominated world of science (and society) is always worth understanding and appreciating. Stein portrays her as a strong minded, generous con- tributor and patron of the sciences. Her grand vision and control of the development of the natural sci- ences at the University of California is only matched by her benevolence, even in spite of those who dis- agreed with her. A remarkable, if not contradictory person. TIM TOKARYK Box 163, Eastend, Saskatchewan SON OTO Canada News and Comment Froglog: Newsletter of the Declining Amphibian Populations Task Force (53) October 2002. Contents: DAPTF Seed Grants 2003 — RANA: The Research & Analysis Network for Neotropical Amphibians — RANA: Red de Analisis sobre Anfibios Neotropicales Amenazados (Bruce E. Young and Karen R. Lips) — Global Amphibian Specialist Group (GASG): Mission and Functions — Publications of Interest — Froglog Shorts. Froglog is the bi-monthly newsletter of the Declining Amphibian Populations Task Force of The World Marine Turtle Newsletter (98) October 2002. Contents: ARTICLES: Preliminary Check- List of the Epizootic Macroalgae Growing on Loggerhead Turtles in the Western Mediterranean Sea —- Recent Sightings of the Green Turtle Chelonia mydas on the Coast of Andhra Pradesh, India — Cayman Turtle Farm Head- starting Project Yields Tangible Success; NoTEs: Results of Experimental Carapace Tagging — Observation of Accidental Capture of a Loggerhead Turtle by Hand-line Fishing in Kefalonia, Greece — Green Turtle Encrusted with White Matter Observed Nesting at Akyatan Beach, Turkey; MEETING REPORTS; BOOK REVIEW; ANNOUNCE- MENTS; News & LEGAL BRIEFS; RECENT PUBLICATIONS. Point Pelee Natural History News 2(3) The fall 2002 issue contains: ARTICLES: Turtle Research at Point Pelee National Park (Connie L. Browne) — Note- worthy Bird Records: June to August 2002 (Alan Worm- ington) — Brown Pelican: New to Point Pelee (Peter S. Burke) — Point Pelee Butterfly Count: August 10, 2002 (Dan Dufour) — Late-Season Butterflies at Point Pelee: Why Here? (Alan Wormington) — IN THE FIELD — NEws AND ANNOUNCEMENTS — UPCOMING EVENTS AND OUTINGS. This newsletter for Point Pelee National Park, Ontario, is edited by Alan Wormington (e-mail: wormington@ Conservation Union (IUCN)/Species Survival Commission (SSC) and is supported by The Open University, The World Congress of Herpetology, and Harvard University. The newsletter is edited by John W. Wilkinson, Department of Biological Sciences, The Open University, Walton Hall, Milton Keynes, MK7 6AA, United Kingdom; e-mail: daptf@open.ac.uk. Funding for Froglog is underwritten by the Detroit Zoological Institute, P. O. Box 39, Royal Oak, Michigan 48068-0039, USA. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Research Group, School of Biological Sciences, University of Wales, Swansea, SA2 8PP Wales, United Kingdom; e-mail MTN @swan.ac.uk; Fax +44 1792 295447. Subscriptions to the MTN and donations towards the production of MTN and its Spanish edition NTM [Noticiero de Tortugas Marinas] should be sent to Marine Turtle Newsletter c/o Chelonian Research Foundation, 168 Goodrich Street, Lunenburg, Massachusetts 01462 USA; e-mail RhodinCRF @ aol.com; fax + 1 978 582 6279. MTN website is: . juno.com). Editorial Assistants are M. Lea Martell and Matthew J. Smith. The web site is www.wincom.net/ ~fopp/Natural_History_News.htm. Subscription rates are Canada: CAN $15 (one year) or $30 (two years); International: US $15 (one year) or $30 (two years). Send payment (and e-mail address, optional) to The Friends of Point Pelee, 1118 Point Pelee Drive, Leamington, Ontario N8H 3V4 Canada. Issues are mailed in March, June, September, and December, and back issues are available for $15 per Volume/ $5 per issue (postage paid). Recovery: An Endangered Species Newsletter (22) The October 2002 issue contains: RECOVERY HIGHLIGHTS: Plovers return home; Reptile recovery; NEWS Bites: Marmot rebounding; COSEWIC Update; CITES Update; RENEW Update; ESRF update; PRoFILe: National parks protect species at risk (Leah de Forest); SPECIAL REPORT: New fund established (Recovering B.C. Plants; Restoring frog habitat; Recovering the Piping Plover; Sydenham River recovery; FiELD Notes: Translocating badgers (Ian Adams); Breeding Wolverines (Sylvie Bouchard); New teams established; ANNOUNCEMENTS: Awards; New publications; Correction; Upcoming events; Site seeing; FEATURED SPECIES: Protecting endangered fish (Mike Pearson). Canadian Wildlife Service, Environment Canada, Ottawa, Ontario K1A 0H3. Web site: www.cws-scf.ec.gc.ca/es/ recovery/archive.html. 680 2002 THE OTTAWA FIELD-NATURALISTS’ CLUB AWARDS 681 National Recovery Plan (23) Long’s Braya (Braya longii Fernald) and Fernald’s Braya (Braya fernaldii Abbe) The Recovery of Nationally Endangered Wildlife [Canada] has issued its “National Recovery Plan for the Long's Braya (Braya longii Fernald) and Fernald’s Braya (Braya fernaldii Abbe)” as Number 23, September 2002. 33 pages. It was prepared by Marilyn F. E. Anions on behalf of the Braya Recovery Team. Copies are available from Recovery Secretariat, c/o Canadian Wildlife Service, Environment Canada, Ottawa, Ontario K1A 0H3; Tel 819- 953-1410; Fax 819-994-3684. E-mail: RENEW- RESCAPE @ec.gc.ca. Response Statements for Extirpated, Endangered and Threatened Species Listed by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) in 2001 This 65-page booklet documents existing conservation measures that federal, provincial, and territorial jurisdic- tions have in place to protect and/or recover 32 species or populations of species of fish (6), amphibians (4), reptiles (3), birds (4), mammals (3), molluscs (2), mosses (2), vas- cular plants (8) newly assessed in 2001 by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) as extirpated (1), endangered (13), or threat- ened (18). Copies are available from Recovery Secretariat RENEW- RESCAPE @ec.gc.ca. See http://www.speciesatrisk.gc.ca/ sar/effors/index.htm. Recovery of Nationally Endangered Wildlife 2001-2002 Annual Report Contents: Report from Co-Chairs — Highlights of 2001-2002 [Of 118 Endangered, 94 Threatened, and 17 Extirpated species on the November 2001 COSEWIC list: 83 have recovery teams totalling 468 team members, 14 have final recovery strategies or plans, 68 have recovery plans or strategies in development, 85 are the direct focus of recovery efforts, 42 others are included in ecosystem recovery, 22 show stable or increasing population trends, $26.2 million expended on recovery (salaries + expenses), employment equivalent to about 158 salaried and 18 vol- unteer people working full-time, and 196 organizations have made financial contributions; 167 public consulta- tion sessions were organized by recovery teams, 22 recov- ery teams are incorporating Aboriginal Traditional Knowledge or community knowledge into recovery plan- ning] — Stewardship and Recovery [web portal www.stewardshipcanada.ca] — Status of recovery (sum- maries for 93 species, 6 ecosystems and 2 Conservation Programs or Landscape Scale Initiatives Benefiting — Financial contributors — Funding per Species or Group — Canadian Wildlife Directors Committee. The report was produced by the Canadian Wildlife Service in cooper- ation with the provinces, territories, Fisheries and Oceans Canada, Parks Canada Agency, authorized Wildlife Management Boards and many individuals and non- government groups. The Ottawa Field-Naturalists’ Club Awards for 2001 At the Club’s Annual Soirée, held on 26 April 2002, at the Unitarian Church of Ottawa, awards were once again given to members, and non-members, who have distinguished themselves by accomplishments in the field of natural his- Honorary Member — Dr. John B. Theberge This honorary membership acknowledges achievements going back more than 30 years. Dr. John Theberge has made outstanding contributions to Canadian natural sci- ences and to the protection of important Canadian natural landscapes. Beyond that, however, he has also effectively brought to the public eye many important natural history issues through numerous books and articles via television and radio, and through a series of engaging and effective speeches from coast to coast. Dr. Theberge is a world authority on the ecology of wolves. With the unflagging and expert collaboration of his wife Mary Theberge, John has spent decades developing a substantial portion of the scientific underpinning for the conservation management of wolves in North America. He has particularly focused his work on the wolves of Ontario’s Algonquin Park, a remarkable population vari- ously described as a unique form of Timber Wolf, as the last truly wild element of the endangered Red Wolf or even as a distinct species. John’s studies of the relationships between wolves and their environments in Algonquin Park, in the Canadian north and elsewhere, have provided chal- lenging research opportunities for dozens of students. A number of these students have gone from the solid scientif- ic and inspirational foundation provided by Dr. Theberge’s academic leadership to develop successful careers in Can- adian biological sciences. Conservation Award, Member — David White The quest to preserve ecologically sensitive lands is made that much stronger when supported by convincing scientific data, including area inventories of fauna and flora. David White has committed much of his time, energy and expertise over many years to collecting such data, greatly contributing to our knowledge of this area’s ecological diversity. David’s documentation of the special botanical features of such areas as the Constance Bay sand dunes, Purdon Fen and Innis Point alvars, all well-known to Ottawa naturalists, have supported conservation protection efforts in these important habitats. His work has also led to ANSI (Areas of Natural and Scientific Interest) designations at Constance Bay and the Burnt Lands. David’s less well known, but equally important, on-going studies of the vascular flora of Lanark County have been pivotal in conservation efforts in that county. David has become the floristic authority for this ecologi- cally complex but poorly documented area of Ontario. He now has documented the presence of approximately 1200 plant species in the county, and the Lanark list has tory and conservation, or by extraordinary activity within the Club. The following citations, a few somewhat abbrevi- ated, were read to the members and guests assembled for the event. Dr. Theberge is also well-known for his contributions to park establishment. His careful scientific documentation, effective political lobbying and eloquent persuasion were highly influential in the protection of several globally important natural areas in Canada’s north, most notably Kluane National Park in Yukon. His technically accurate and at times lyrical book “Kluane, Pinnacle of the Yukon” attests to his understanding that public awareness and “buy- in’ are important elements of such conservation initiatives. And this conservation focus does not apply only to northern areas. That is demonstrated by his editorship of the beauti- ful and authoritative Legacy: the Natural History of Ontario, as well as numerous contributions to scientific journals across North America, including The Canadian Field-Naturalist. Dr. Theberge was a professor of ecology and environmen- tal studies at the University of Waterloo from 1972 to 2000. He has received many conservation awards during his career. At the close of his speech accepting the Harkin Conservation Award from the Canadian Parks and Wilderness Society, he led the audience in a group wolf howl, saying, “This is a bonding for all of us who work for wild places.” He has demonstrated mastery in both the sciences and the arts and has applied these exceptional skills to a greater purpose: the preservation of biodiversity. The natural and wild landscapes of Canada will benefit indefinitely from this exceptional record of scientific and conservation achievements. become a key component of a province-wide floristic atlas. David’s floristic work is providing valuable resource information for various local conservation initia- tives such as conservation/protection arguments for vari- ous natural environment assessments, for example, the Highway 7 upgrade planning and highway service road design studies in the Carleton Place area by various envi- ronmental consultants. David has published dozens of articles on botanically interesting and important natural areas, including many contributions to Trail & Landscape. He is also co-author of the well-known and much-used 1978 Checklist of Vascular Plants of the Ottawa-Hull Region, Canada, and the impor- tant 1993 publication, Invasive Plants of Natural Habitats of Canada. We are pleased to acknowledge the many contributions David White has made to the preservation of natural areas and to the further understanding of our botanical legacy, by presenting him with the 2001 Conservation Award for OFNC members. 682 2002 THE OTTAWA FIELD-NATURALISTS’ CLUB AWARDS 683 Conservation Award, Non-Member — Land Preservation Society of the Ottawa Valley (President Marc Stabb accepted the award on behalf of the Society) The Conservation Award for a non-member to the Land Preservation Society of the Ottawa Valley for its hard-won victory in saving Gillies Grove, a magnificent old-growth forest in Arnprior, from falling victim to a housing devel- opment. The community’s struggle to save Gillies Grove was taken up by the Society although it faced the formidable challenge of raising half a million dollars in a small local community for the outright purchase of the for- est. It took several years of unrelenting effort, before this goal was finally realized. The focus of the struggle then changed to one of negotiating to balance several conflicting interests: the real estate ambitions of the landowner, the revenue base of the town of Arnprior, and the need to impose land-use restrictions that would safeguard the integrity of the old-growth forest. Year after year, negotia- George McGee Service Award — Fenja Brodo It is hard to adequately describe and acknowledge the all around excellent manner in which Fenja Brodo managed and directed the production of almost ten volumes of Trail & Landscape throughout her editorship. Not content with simply waiting for manuscripts to arrive, Fenja asked many amateurs and professionals to share their observations and insights on a wide variety of natural history topics. She par- ticularly encouraged the contributions of young people and inexperienced authors, and she enthusiastically sought out the best natural history poets to add dimension to the jour- nal. Her keen interest in natural history and scientific back- ground led her not only to edit manuscripts, but also to test tions faltered as new objections or new obstacles were raised, dashing the hopes of the Society and the communi- ty. The Land Preservation Society refused to give up, adamant in its resolve that this old-growth forest must not be sacrificed to development. After an epic eleven-year struggle, the unswerving commitment of the Land Preservation Society of the Ottawa Valley resulted in the purchase of Gillies Grove by the Nature Conservancy of Canada in December of 2001, with stewardship of the for- est undertaken by the local Macnamara Field Naturalists’ Club. From all of us in the Ottawa Field-Naturalists’ Club to the Land Preservation Society of the Ottawa Valley, we say, “Well done” and “Thank you!” You are most deserv- ing of our Conservation Award for 2001. keys in the field and meticulously check facts against reli- able literature. Many authors thanked her for her attention to detail, and for her patience and diplomacy. Always sen- sitive to the needs of the Club, she carefully prepared the publication team for her replacement so that the transition was seamless for the reader and for the hard working pro- duction group as well. In fact, she continues to serve as an associate editor, helping our new editor, Karen McLachlan Hamilton, whenever she is asked, and carefully proofread- ing each issue. Fenja’s dedicated service as editor of T&L for most of the 1990s makes her eminently qualified for the Club’s service award. Anne Hanes Natural History Award — Robert Bracken and Christina Lewis Some animals seem to get overlooked by naturalists despite being colourful, lively, and diverse. Dragonflies and damselflies are good examples. Fortunately for OFNC members, Bob Bracken and Chris Lewis remedied this sit- uation by making these insects the object of their keen observational skills over the past five years. Their passion for “dragons,” as they call them, is demonstrated by their remarkable checklist of the species found in the Ottawa Region and published in Trail & Landscape in 1998. The list is packed with information on the precise habitat pref- erences of each species, their flight periods, and notes on their abundance in the region. It is based on a great deal of field work by Bob and Chris all over the region, supple- mented by careful attention to the scientific literature. As Member of the Year — Eve Ticknor In honouring Eve Ticknor as Member of the Year, we do indeed mark a year of tremendous and devoted service to the club. Many club members will know Eve as the OFNC’s Peregrine Falcon watch coordinator for the last three years. This year-round job, all voluntary, requires stamina, patience and diplomacy. For several weeks every year, it is intense and sometimes heartbreaking. There are few, if any, other volunteer jobs within the OFNC, with the Chris and Bob continued their “quest for dragons” (to use their own words), they frequently visited Morris Island, which turns out to be a Mecca for dragonfly and damselfly enthusiasts. To share their deep interest in these fascinat- ing insects, Bob and Chris led a field trip for the OFNC in July of 2000 finding 30 of the 66 species they have record- ed so far from the island. All their exciting finds were recorded in a fine article in T&L, illustrated by beautiful and accurate drawings by Chris. For their enthusiastic work in opening the world of Odonates (“dragons”) to members of the club and publishing two fine articles on their natural history, the Club is proud to present the Anne Hanes Natural History Award to Robert Bracken and Christina Lewis. possible exception of gardening, where one deals with mat- ters of life and death. Every year the falcon watchers begin the breeding season full of hope: that the peregrines will nest, that young birds will hatch, that fledglings will clam- our, then soar, safely amidst the downtown towers, One thing that makes Eve such a good falcon watch coordinator is that she loves the falcons, 2001 was a partic- ularly trying year for Eve and the other volunteers who 684 dedicate themselves to ensuring the birds’ survival. In 2001, peregrine parents Connor and Horizon produced three eggs. One hatched but the other two didn’t. When eggs turn out not to be viable, it’s possible to adopt. In 2001 the Canadian Peregrine Foundation purchased a hatchling to accompany the bird that did hatch. The parents fed and nurtured the young birds — chicklets, as Eve would call them — until they were ready to fly. In years past, young falcons, and even once an adult, have flown to their death by crashing into glass-walled buildings, thinking the reflections ahead were merely open sky. Every year, falcon watchers and others have tried to find ways to break up the reflections. In 2001, with the assistance and leadership of the Ontario Ministry of Natural Resources, the most dangerous nearby building was fes- tooned with colourful flags to make it look more like the imposing wall that it was. In spite of this effort, the 2001 watch was cut short. On June 24th, on their first day of flying, both young birds died when they were unable to fly strongly enough to avoid col- liding with a different building. The falcon watch goes on, every year with new hope. Every year requires building and rebuilding relationships with downtown property owners, building managers, the Ontario THE CANADIAN FIELD-NATURALIST ,, — eee on ar a nn cee irre Vol. 116 “ Ministry of Natural Resources, the Canadian Peregrine Foundation, many volunteers and members of the public. Eve further distinguished herself in 2001 by taking on the job of bird feeder liaison for the birds committee. This much needed role involves fielding comments and concerns from members and the public, and directing them to the appropriate bird feeder caretaker. It requires tact, problem- solving skills and energy. Eve’s willingness to help here has been a great blessing. Other falcons stole Eve’s heart in 2001, too. She indepen- dently conducted a survey of urban-nesting Merlins in Ottawa. She has jumped in to assist with other projects, including helping with local arrangements for the annual Taverner Cup birding competition. We’re sure there must be more than we know about. Eve tends to take on work quietly, just pushing up her sleeves and doing it. Eve is also active in her housing co-op, including as a choir member. Somehow she holds down a day job as a teaching assistant for special- needs children. Eve is busy every year, but we recognize 2001 as a special year of service and achievement. IRWIN M. BRODO, CHRISTINE HANRAHAN, BEVERLY MACBRIDE, and SHEILA THOMSON CFNC Awards Committee Index to Volume 116 Compiled by Leslie Durocher Abies balsamea, 44,70,317,476,524,572,588,602,641 Abraham, K. F., 42 Accipiter cooperii, 137,580 Accipiter cooperii, Mobbing Black-billed Magpie, Pica hudsonia, Killed by Cooper’s Hawk, 137 Accipiter cooperii, Successfully Nest at High Densities in the Northern Great Plains, Cooper’s Hawks, 580 Acer pensylvanicum, 525 rubrum, 108,317,476,525,530,588 saccharum, 476,524,588 spicatum, 525,600 Achillea millefolium, 55 millefolium ssp. lanulosa, 446 sibirica, 196,446 Acipenser fulvescens, 434 oxyrhynchus, 434 Acorn, 109 Adams, B. W., 51 Aechmophorus occidentalis, 366 Agoseris glauca, 55,198 Agoseris, Pale, 293 Agosta, S. J., K. M. Kuhn, and D. Morton. Bat Night Roost at an Abandoned Mine in Western Maryland, 389 Agropyron spp., 52 dasystachyum, 53 smithii, 53 spicatum, 58 subsecundum, 53 Agrostis spp., 482 aequivalvis, 216 borealis, 210 hyemalis, 575 mertensii, 199 thurberiana, 199 Aiken, S. G. and R. A. Buck. Aquatic Leaves and Regener- ation of Last Year’s Straw in the Arctic Grass, Arctophila fulva, 81 Aix sponsa, 45 Alaska, Diets of Northern Flying Squirrels, Glaucomys sabrinus, in Southeast, 98 Alaska in Wrangell-St. Elias National Park and Preserve with Comments on the Floristics, Notable Vascular Plants from, 192 Alaska, Predation by Wolves, Canis lupus, on Wolverines, Gulo gulo, and an American Marten, Martes ameri- cana, in, 132 Alaskan Material of Native and Naturalized Mustards, Brassicaceae (Cruciferae), Chromosome Numbers Determined from Canadian and, 611 Alberta Alfalfa Fields, Reproductive Characteristics of Northern Pocket Gophers, Thomomys talpoides, in, 319 Alberta using Call Playbacks, Night Surveys of Yellow Rails, Coturnicops noveboracensis, and Virginia Rails, Rallus limicola, in, 408 Alberta Wildlife Status Reports, 178,356 Alberta, Winter Habitat Selection at Three Spatial Scales by American Elk, Cervus elaphus, in West-Central, 183 Alces alces, 125,127,132,142,417,488 Alder, 70 Green, 184 Speckled, 602 Alewife, 359 Alfalfa, 319,322 Alfalfa Fields, Reproductive Characteristics of Northern Pocket Gophers, Thomomys talpoides, in Alberta, 319 Alisma plantago-aquatica, 566 Alle alle, 406 Alliaire officinale, 623 Alliaria officinalis, 612,623 petiolata, 612,623 Allium schoenoprasum ssp. sibiricum, 446 tricoccum, 114 Allysson maritime, 628 Alnus sp., 70 crispa, 184 incana, 588 rugosa, 525 Alopecurus alpinus, 199 borealis, 210 pratensis, 448 Alopex lagopus, 22 Alosa pseud oharengus, 359 Alvo, R., C. Blomme and D. V. Weseloh. Double-crested Cormorants, Phalacrocorax auritus, at Inland Lakes North of Lake Huron, Ontario, 359 Alysson a calices persistants, 623 de déserts, 623 Alyssum alyssoides, 612,623 americanum, 612 desertorum, 612,623 murale, 624 obovatum, 612 saxitale, 624 Alyssum, Hoary, 624 Alyssum, Small, 623 Alyssum, Sweet, 628 Alyssum, Yellow, 623 Ambloplites rupestris, 434 Ambystoma laterale, 554 maculatum, 554 Ameiurus nebulosus, 434 Amelanchier sp., 525,590 alnifolia, 446 Amerorchis rotundifolia, 446 Anabaena azollae, 441 Anaphalis margaritacea, 525 Anarhichas denticulatus, 423 lupus, 423 minor, 423 Anarhichas lupus, in Canada, A Review of the Status of the Atlantic Wolffish, 423 Anas spp., 368 acuta, 45,368,404 685 686 americana, 45,368,404 clypeata, 45 crecca, 45 discors, 45 platyrhynchos, 45,368 rubripes, 44 Andersen, J. J. Status of Redside Dace, Clinostomus elon- gatus, in the Lynde and Pringle Creek Watersheds of Lake Ontario, 76 Anderson, S. H., 60 Androsace alaskana, 282 chamaejasme ssp. lehmanniana, 446 septentrionalis, 55 Anemone multifida, 55 Anemone, Plume, 460 Anser albifrons, 403 Antelope, Pronghorn, 142 Antennaria media, 199 monocephala ssp. monocephala, 446 pulcherrima, 446 Anthus rubescens, 407 Antilocapra americana, 12,142 Apalone spinifera, 104,554 Apalone spinifera, Nesting Activities of an Eastern Spiny Softshell Turtle, 104 Aphragmus eschscholtzianus, 199 Aphriza virgata, 366 Apple, 92 Crab, 306 Apple Trees, Malus pumila, in Northeast North America, Black Bear, Ursus americanus, Hair and, 305 Applegate, R. D., Reviews by, 166,334 Aquilegia canadensis, 575 Arabette des dames, 624 glabre, 624 hirsute, 624 Arabidopsis lyrata ssp. kamchatica, 613 lyrata ssp. lyrata, 613 salsuginea, 612 thaliana, 612,624 Arabis albida, 612 alpina ssp. caucasica, 612 arenicola, 612 arenicola var. pubescens, 254 calderi, 197 caucasica, 612,624 codyi, 198 columbiana, 446,612 divaricarpa, 612 divaricarpa var. dacotica, 612 divaricarpa var. divaricarpa, 612 drepanoloba, 199 drummondii, 612 eschscholtziana, 612 exilis, 612 glabra, 613,624 hirsuta, 613 hirsuta ssp. eschscholtziana, 612 hirsuta var. hirsuta, 624 hirsuta var. pycnocarpa, 613 holboellii, 612 holboellii var. consanguinea, 613 holboellii var. pendulocarpa, 612 holboellii var. retrofracta, 613 holboellii var. secunda, 446,613 THE CANADIAN FIELD-NATURALIST Vol. 116 e kamtschatica, 613 lemmonii, 197,613 lemmonii var. drepanoloba, 250 lyallii, 250,613 lyrata, 613 lyrata ssp. kamchatica, 613 media, 199,613 microphylla, 613 nuttallii, 446 pendulocarpa, 612 petraea, 613 pinetorum, 446,613 sparsiflora var. columbiana, 613 Arabis, Hairy, 624 Aralia hispida, 525 nudicaulis, 70,525,575,600 Archilochus colubris, 487 Arctagrostis latifolia, 83 Arctophila fulva, 81 Arctophila fulva, Aquatic Leaves and Regeneration of Last Year’s Straw in the Arctic Grass, 81 Arctostaphylos rubra, 446 uva-ursi, 446,575,675 Ardea herodias, 359 herodias fannini, 604 Arenaria capillaris, 199 interpres, 405 longipedunculata, 199,446 melanocephala, 366 Armeria maritima ssp. arctica, 446 Armoracia rusticana, 624 Arnica amplexicaulis ssp. prima, 199 latifolia, 199 mollis, 199 Arnica, Broadleaf, 293 Hairy, 293 Arrow-grass, Seaside, 450 Arrowhead, Arum-leaved, 450 Artemisia alaskana, 196,446 arctica, 196 biennis, 448 campestris, 446 frigida, 55 furcata, 293 hyperborea, 199 michauxiana, 446 tilesii, 446 Ash, Black, 598 Green, 580 White, 524,593 Asio flammeus, 406 Aspen, 485 Quaking, 581,641 Trembling, 44,69, 136,137,184,458,524,575,602 Asphodel, Northern False, 457 Aster spp., 482,525,590 acuminatus, 600 alpinus ssp. vierhapperi, 199 borealis, 199 ciliolatus, 575 junciformis, 296 laevis, 55 macrophyllus, 575,641 puniceus, 600 sibiricus, 446 2002 umbellatus, 600 Aster, Alpine, 296 Arctic, 472 Ciliolate, 575 Large-leaved, 575 Astragalus adsurgens var. tananaicus, 272 adsurgens ssp. viciifolius, 199 alpinus, 446 australis, 446 bodinii, 446 canadensis, 641 eucosmus ssp. sealei, 199 harringtonii, 199 neglectus, 575,640 nutzotinensis, 200 pectinatus, 55 robbinsii ssp. harringtonii, 274 robbinsii var. harringtonii, 274 williamsii, 200,446 Astragalus neglectus, in an Eastern Ontario Alvar Wood- land, Initial Beneficial Effects of Fire and Bulldoz- ing on Neglected Milkvetch, 640 Athene cunicularia, 1,307 Athene cunicularia, Found in Saskatchewan, Large Clutch Size of a Burrowing Owl, 307 Athyrium filix-femina, 525,600 Athysanus pusillus, 614 Atkinson, J. E., Review by, 677 Avens, Large-leaved, 465 Yellow, 465 Awlwort, 266 Aythya affinis, 45,368 collaris, 45 marila, 366 Azolla caroliniana, 441 filiculoides, 441 mexicana, 441 pinnata, 441 Azolla caroliniana, at Ottawa, Ontario, Ephemeral Occur- rence of the Mosquito Fern, 441 Badger, 307 Barbarea orthoceras, 614 stricta, 624 verna, 624 vulgaris, 614,624 Barbarée printaniére, 624 raide, 624 vulgaire, 624 Barley, Foxtail, 453 Barry, R. E., 529 Basil, Wild, 575 Bass, Rockbar, 434 Smallmouth, 359,434,539 Striped, 434 White, 35,434 Bat, Big Brown, 389,645 Hoary, 124,136 Little Brown, 389 Red, 390 Silver Haired, 646 Small-footed, 390 Bat, Eptesicus fuscus, in New Brunswick, Over-wintering and Reproduction by the Big Brown, 645 Bat, Lasiurus cinereus (Chiroptera: Vespertilionidae), from INDEX TO VOLUME 116 687 Prince Edward Island, First record of the Hoary, 124 Bat Night Roost at an Abandoned Mine in Western Maryland, 389 Bats, Lasiurus cinereus: A Cautionary Note, Apparent Capture Myopathy in Hoary, 136 Bean, Buffalo, 54 Bear, Black, 63,125,132,305,313, 418 Brown, 418 Grizzly, 63,132,313 Polar, 22,313,324 Bear, Ursus americanus, Hair and Apple Trees, Malus pumila, in Northeast North America, Black, 305 Bearberry, 575 Common, 469 Red Alpine, 469 Beardtongue, Gorman’s, 292 Beaudoin, C., 523 Beaver, 60,118,417,648 Beavers, Castor canadensis, in Wyoming, Survival, Fates, and Success of Transplanted, 60 Beckmannia syzigachne, 566 Beech, 485 American, 306,593,602 Bellflower, Yukon, 471 Bennett, B., 446 Bentgrass, Northern, 216 Red, 210 Thurber’s, 210 Berteroa incana, 614,624 Bertéroa blanc, 624 Betula alleghaniensis, 317,476,524,588,602 cordifolia, 476 glandulosa, 446 lenta, 530 neoalaskana, 446 papyrifera, 44,70,137,317,476,524,588,602 populifolia, 317,525 Bigeye, 632 Short, 632 Bindweed, Low, 577 Biodiversity: Journal of Life on Earth 3(1) February 2002, i77 Birch, Alaska, 459 Black, 530 Ground, 459 Mountain, 476 White, 44,70, 137,476,524,602 Yellow, 476,524,593 Bison bison, 141,313 bison athabascae, 157 bonasus, 147 Bison, 313 Wood, 146 Bison bison: A Tale of Two Herds, Canada and the “Buffalo”, 141 Bistort, Alpine, 459 Bitter-cress, Alpine, 461 Cuckoo, 462 Blackberry, 530 Bladderpod, Arctic, 264,463 Bladderwort, Greater, 470 Blomme, C., 359 Blue Bell, 470,575 Blueberry, 99,530 688 Bluebird, Eastern, 479 Bluefish, 134 Boechera divaricarpa, 612 drummondii, 612 Bonasa umbellus, 2,487 Book-review Editor’s Report for Volume 115 (2002), 358 Boreal Dip Net: Newsletter of the Canadian Amphibian and Reptile Conservation Network, The, 177,355 Bork, E. W., B. W. Adams, and W. D. Willms. Resilience of Foothills Rough Fescue, Festuca campestris, Rangeland to Wildfire, 51 Bos mutus, 143 Boschniakia rossica, 196 Botrychium ascendens, 200 boreale, 202 lanceolatum, 200 minganense, 200 pinnatum, 200 Bouleau blanc, 317 gris, 317 jaune, 317 Bourse-a-pasteur, 625 Bowman, J., Reviews by, 344,346 Boyd, W. S., 366 Boykinia richardsonii, 446 Boykinia, Richardson’s, 465 Brachyramphus marmoratus, 366 Bracken-fern, Eastern, 578 Brant, 366,397 Branta bernicla, 84,366 bernicla nigricans, 403 canadensis, 44,368,403 Brassica campestris, 448,614,625 juncea, 614,624 napus, 625 nigra, 625 oleraceae, 625 rapa, 461,614,623 Braya glabella, 614 glabella ssp. glabella, 200 glabella ssp. purpurascens, 200 humilis, 614 purpurascens, 254,614 thorild-wulfii, 614 Braya, 254 Purplish, 254 Breton, L., 488 British Columbia, Population Trends of Nesting Glaucous- winged Gulls, Larus glaucescens, in the Southern Strait of Georgia, 603 British Columbia, Seasonal Distribution of Waterbirds in Relation to Spawning Pacific Herring, Clupea pal- lasi, in the Strait of Georgia, 366 British Columbia’s Central and North Coast: Distribution and Conservation Assessment, The Gray Wolves, Canis lupus, of, 416 British Isles and Eastern Canada, North Atlantic Sperm Whale, Physeter macrocephalus, Strandings on the Coastlines of the, 371 Broders, H. G., 645 Brome, Kalm’s, 575 Pumpelly, 451 Smooth, 451 Bromus inermis, 448 kalmii, 575 THE CANADIAN FIELD-NATURALIST ae Se EDD A ie ae ya e Vol. 116 a pumpellianus var. arcticus, 446 Brown, G. S., 122 Bryoria spp., 102 Bubo virginianus, 138,582 Bucephala albeola, 45,366 clangula, 45,366 islandica, 366 Buck, R. A., 81 Buckbean, 469 Buckthorn, Glossy, 575 Budworm, Spruce, 476 Buffalo, 141 Black, 434 Buffalo, Bison bison: A Tale of Two Herds, Canada and the, 141 Buffalo-berry, Canada, 184 Bufflehead, 45,366 Bufo americanus, 552 boreas, 117 hemiophrys, 117 Bulleye, 632 Bullfrog, 554 Bullhead, Brown, 434 Bunchberry, 70,468 Bunias orientalis, 625 Bunias d’ Orient, 625 Bunting, Snow, 397 Bur-reed, 450,566 Arctic, 207 Small, 450 Burbot, 39 Burnet, Great, 466 Bush-cranberry, Low, 471 Buteo jamaicensis, 137 lagopus, 404 Butler, R.W., 366 Buttercup, Northern, 248 Northern Seaside, 460 Pacific, 248 Snow, 461 Sulphur, 248 Cabbage, Deer, 287 Skunk, 99 Steppe, 629 Cactus, Brittle Prickly-pear, 547 Kaladar, 547 Cactus (Opuntia fragilis), Bill Dore’s Notes on the Kaladar, 547 Cairns, D: K., 22 Cakile edentula var. edentula, 614,625 edentula var. lacustris, 614 maritima, 614,625 Calamagrostis canadensis, 184 canadensis ssp. langsdorfii, 446 stricta ssp. stricta, 447 Calcarius lapponicus, 407 Calidris alba, 405 alpina, 405 bairdii, 405 fuscicollis, 405 maritima, 405 mauri, 405 melanotos, 405 minutilla, 405 2002 pusilla, 405 Calla palustris, 447 Calla, Wild, 456 Callitriche sp., 566 anceps, 200 hermaphroditica, 200 heterophylla ssp. heterophylla, 278 Caltha leptosepala, 200 Calypso bulbosa, 196,447 Calystegia spithamaea, 575 Camelina alyssum, 625 microcarpa, 614,625 parodii, 625 sativa, 614,625 Caméline a petits fruits, 625 alysson, 625 cultivée, 625 Campanula aurita, 447 rotundifolia, 575 uniflora, 447 Campion, Menzies’, 242 Moss, 460 Williams’, 242 Canada, A Review of the Status of the Atlantic Wolffish, Anarhichas lupus, in, 423 Canada and the “Buffalo”, Bison bison: A Tale of Two Herds, 141 Canada, First Report of the Rare Charophyte Nitella macounii (T. F. Allen) T. F. Allen in Saskatchewan and Western, 559 Canada, Narwhal, Monodon monoceros, Near Western Baffin Island, Nunavut, 323 Canada, North Atlantic Sperm Whale, Physeter macro- cephalus, Strandings on the Coastlines of the British Isles and Eastern, 371 Canada, Weedy Introduced Mustards (Brassicaceae) of, 623 Canadian and Alaskan Material of Native and Naturalized Mustards, Brassicaceae (Cruciferae), Chromosome Numbers Determined from, 611 Canadian Orchidologist, A Tribute to Edward Warren Greenwood (1918-2002), 326 Canadian Prairies and adjacent areas, Spread and Disap- pearance of the Greater Prairie-Chicken, Tympanu- chus cupido, on the, 1 Canadian Species at Risk May 2002, 512 Candytuft, False, 266 Globe, 627 Rocket, 627 Canis familiaris, 125,418 latrans, 63,127,129, 132,310,418,649 lupus, 125,127,132,139,148,311,313,315,416,648 Canis familiaris, Activities in Central Yukon, Wolf, Canis lupus, Response to Domestic Sled Dog, 125 Canis latrans, and Red Fox, Vulpes vulpes, in Ontario: Implications for Disease-spread, Long Distance Movement by a Coyote, 129 Canis latrans, in Labrador, First Record of an Eastern Coyote, 127 Canis lupus, from Chest Girth Measurements, Estimating the Weight of Wolves, 313 Canis lupus, in Relation to Latitude, Breeding Season of Wolves, 139 Canis lupus, of British Columbia's Central and North Coast: Distribution and Conservation Assessment, INDEX TO VOLUME 116 689 The Gray Wolves, 416 Canis lupus, on Wolverines, Gulo gulo, and an American Marten, Martes americana, in Alaska, Predation by Wolves, 132 Canis lupus, Pack Territory Edge and Core, Differential Use of a Wolf, 315 Canis lupus, Response to Domestic Sled Dog, Canis famil- iaris, Activities in Central Yukon, Wolf, 125 Canis lupus, Suffering from Sarcoptic Mange: Importance of Sequential Monitoring, Behavioral Modification of Gray Wolves, 648 Canis lupus, Territorial Marking by Lone Male Gray Wolves, 311 Capsella bursa-pastoris, 448,614,625 Caquillier édentulé, 625 maritime, 625 Carassius auratus, 434 Cardamine angulata, 614 bellidifolia, 196,447,615 cocatenata, 615 digitata, 447,615 diphylla, 615 flexuosa, 625 hirsuta, 615,625 impatiens, 625 microphylla, 615 microphylla ssp. blaisdellii, 615 occidentalis, 615 oligosperma, 615 parviflora var. arenicola, 615 pensylvanica, 615 pratensis, 447,615,626 purpurea, 615 umbellata, 615 Cardamine des prés, 626 flexueuse, 625 hirsute, 625 impatiente, 625 Cardaria chalepensis, 615,626 draba, 615,626 pubescens, 615,626 Carduelis flammea, 407 pinus, 73 Carex spp., 54,581,590 adelostoma, 200 albo-nigra, 200 aquatilis ssp. aquatilis, 447 aquatilis var. stans, 396 bebbi, 575 buxbaumii, 200 capillaris, 196 castanea, 575 chordorrhiza, 200 concinna, 447 crawfordii, 200 eburnea, 200,575 enanderi, 224 filifolia, 200 gynocrates, 447 holostoma, 200 houghtoniana, 575 interior, 200 krausei, 200 lachenallii, 196 lasiocarpa ssp. americana, 200,447 690 laxa, 195 lenticularis var. dolia, 200 leptalea, 200 limosa, 196 lugens, 447 macrocheata, 196 misandra, 447 nardina, 196,447 nigricans, 200 obtusata, 200 oederi ssp. viridula, 228 parryana, 198 pauciflora, 200 pellita, 447 petasata, 197 petricosa, 200 phaeocephala, 200 podocarpa, 196,447 praticola, 200 richardsonii, 575 rupestris, 447 saxatilis, 447 scirpoidea, 447 spectabilis, 447 stylosa, 200 subspathacea, 81 tenuiflora, 196,447 vaginata, 447 viridula, 200 williamsii, 200 Caribou, 122,127,313,400,488 Carp, Common, 30,434 Carpiodes carpio, 30 Carpodacus purpureus, 487 Carpsucker, River, 30 Cassiope mertensiana, 200 tetragona, 396 Castilleja, 195 chrymactis, 200 elegans, 200 unalaschensis, 197 yukonis, 200 Castor canadensis, 60,118,417,648 fiber, 65 Castor canadensis, in Wyoming, Survival, Fates, and Suc- cess of Transplanted Beavers, 60 Casuarina equisetifolia, 636 Catchfly, Arctic, 240 Largefruit, 242 Catharus guttatus, 487 ustulatus, 487 Catling, P. M., 112 Catling, P. M. and A. Sinclair. Initial Beneficial Effects of Fire and Bulldozing on Neglected Milkvetch, Astragalus neglectus, in an Eastern Ontario Alvar Woodland, 640 Catling, P. M., A. Sinclair, and D. Cuddy. Plant Com- munity Composition and Relationships of Disturbed and Undisturbed Alvar Woodland, 571 Catostomus catostomus, 434 commersoni, 79,434,537 Catseye, Shacklett’s, 291 Cattail, 442 Common, 566 Cedar, Eastern White, 122 THE CANADIAN FIELD-NATURALIST Vol. 116 s Northern White, 641 Red, 99 White, 525,593 Yellow, 99 Centaurea nigra, 525 Centrocercus urophasianus, | Cepphus columba, 368 grylle, 406 Cerastium arvense, 55 fontanum, 575 regelii, 195 Ceratophyllum demersum, 198 Certhia americana, 487 Cervus elaphus, 122,142,183,488 Cervus elaphus, in the French River Delta, Ontario, Ano- malies in Incisor Wear of American Elk, 122 Cervus elaphus, in West-Central Alberta, Winter Habitat Selection at Three Spatial Scales by American Elk, 183 Chaenorrhinum minus, 575 Chagnon, Y., 104 Chamaecyparis nootkatensis, 99 Chamaedaphne calyculata, 196,447,601 Chamaerhodos erecta ssp. nuttallii, 200 Chara braunii, 560 contraria, 568 fragilis, 566 globularis, 566 longifolia, 564 zeylanica, 562 Charadrius semipalmatus, 405 vociferus, 487 Charophyte Nitella macounii (T. F. Allen) T. F. Allen in Saskatchewan and Western Canada, First Report of the Rare, 559 Chelydra serpentina, 106,552 Chen caerulescens, 403 rossii, 403 Chenopodium album, 448 Cherry, Black, 306 Chestnut, 530 Chickadee, Black-capped, 138,479 Boreal, 479 Chickweed, 575 Larger Mouse-ear, 577 Regel’s, 236 Chiogenes hispidula, 525 Chordeiles minor, 487 Chorispora tenella, 626 Chorispora fluet, 626 Choristoneura fumiferana, 476 Chrysanthemum leucanthemum, 575 Chrysemys picta, 552 Chub, Creek, 537 Flathead, 26 Lake, 537 Chubbs, T. E. and F. R. Phillips. First Record of an Eastern Coyote, Canis latrans, in Labrador, 127 Chubs, Platygobio gracilis, in the Upper Missouri River: The Biology of a Species at Risk in an Endangered Habitat, Flathead, 26 - Cicuta douglasii, 280 maculata var. angustifolia, 200 Cinquefoil, Drummond’ s, 272 Marsh, 465 2002 INDEX TO VOLUME 116 691 Norwegian, 465 Pennsylvania, 272 Prairie, 465 Rocky Mountain, 272 Silvery, 575 Tall, 270 Two-Flower, 270 Varileaf, 270 Circea lutetiana, 525 Circus cyaneus, 138,582 Cirsium arvense, 575 vulgare, 575 Citellus variegatus, 530 Cladina, 590 rangiferina, 601 stellaris, 601 Cladothamnus pyrolaeflorus, 200 Clangula hyemalis, 366,404 Claytonia bostockii, 234 tuberosa, 200 Clemmys guttata, 554 insculpta, 555 Clinopodium vulgare, 575 Clinostomus elongatus, 76 Clinostomus elongatus, in the Lynde and Pringle Creek Watersheds of Lake Ontario, Status of Redside Dace, 76 Clintonia borealis, 525,600 Clover, Alsike, 467,579 Red, 467,575 White, 467 Club-moss, Bristly, 448 Common, 448 Mountain, 448 Clupea pallasi, 366 Clupea pallasi, in the Strait of Georgia, British Columbia, Seasonal Distribution of Waterbirds in Relation to Spawning Pacific Herring, 366 Coad, B. W. and J. Gilhen. Additional Records of Bigeye Fishes (Priacanthidae) from the Atlantic Coast of Nova Scotia, Including the First Record of the Glasseye Snapper, Heteropriacanthus cruentatus, 632 Coad, B. W. and J. Gilhen. A Rare Leucistic Spiny Dog- fish, Squalus acanthias, from the Bay of Fundy, Nova Scotia, 120 Cochlearia groenlandica, 615 officinalis, 615 Cod, Atlantic, 424 Cody, W. J., 547 Cody, W. J., Reviews by, 168,344,670 Cody, W. J., C. E. Kennedy, B. Bennett, and V. Loewen. New Records of Vascular Plants in the Yukon Territory IV, 446 Coeloglossum viride ssp. bracteatum, 447 Colaptes auratus, 487 Coe, Fs... 522 Colgan, P., Review by, 345 Collomia linearis, 200 Colpodium vahlianum, 198 Coltsfoot, Sweet, 472 Columbine, Wild, 575 Comandra, Northern, 459 Commandra pallida, 55 Companula rotundifolia, 55 Conringia orientalis, 626 Contopus virens, 487 Cook, F. R., Reviews by, 653,654,655,656,662.663.664, 665 666,667,668 Cook, J. A., 98 Cook, M. B. and C. A. Roland. Notable Vascular Plants from Alaska in Wrangell-St. Elias National Park and Preserve with Comments on the Floristics, 192 Cookeolus boops, 632 japonicus, 632 Coptis groenlandica, 525 trifolia, 200,600 Corallorhiza trifida, 447 Coralroot, Striped, 457 Corbeille d’argent, 624 Coregonus clupeaformis, 125,434 Cormorant, Brandt’s, 366 Double-crested, 359 Pelagic, 366 Cormorants, Phalacrocorax auritus, at Inland Lakes North of Lake Huron, Ontario, Double-crested, 359 Corne-de-cerf didyme, 626 écailleuse, 626 Cornus canadensis, 70,447 ,482,525,601 sericea, 600 stolonifera, 447,525 Coronopus didymus, 626 squamatus, 626 Corvus brachyrhychos, 138 corax, 406 Corydalis aurea ssp. aurea, 575 Corydalis, Golden, 577 Corylus cornuta, 70,525,532 Cotton-grass, Arctic, 230 Narrow-leaved, 455 Sheathed, 455 Tassel, 230 Cottontail, Appalachian, 529 Eastern, 529 New England, 529 Cottontail, Sy/vilagus obscurus, Selection, Size, and Use of Home Range of the Appalachian, 529 Cottonwood, 132 Cottus bairdi, 79 Coturnicops noveboracensis, 408 Coturnicops noveboracensis, and Virginia Rails, Rallus limicola, in Alberta using Call Playbacks, Night Surveys of Yellow Rails, 408 Couesius plumbeus, 537 Cougar, 418 Couture, B., 317 Coyote, 63,129,132,310,418,649 Eastern, 127 Coyote, Canis latrans, and Red Fox, Vulpes vulpes, in Ontario: Implications for Disease-spread, Long Distance Movement by a, 129 Coyote, Canis latrans, in Labrador, First Record of an Eastern, 127 Crab, 428 Crabapple, 92 Cranberry, Bog, 469 Low-bush, 184 Crane, Sandhill, 394 Crane’s-bill, Bicknell’s, 575 Cranson dravier, 626 692 rampant, 626 velu, 626 Crassula aquatica, 446 Crataegus spp., 92 Creeper, Brown, 479 Crepis nana, 447 Cress, Aleutian, 249 Amphibious Water, 629 Asiatic, 629 Austrian Yellow, 629 Creeping Wart, 626 Creeping Yellow, 630 Early Winter, 624 Garden, 628 Globe-podded Hoary, 626 Heart-podded Hoary, 626 Hoary Bitter, 625 Hoary Yellow, 464 Lens-podded Hoary, 626 Lesser Swine, 626 Mitre, 629 Narrow Leaved Bitter, 625 Penny, 464 Shepherd’s, 630 Small Flowered Winter, 624 Wavy Bitter, 625 Cresson amphibie, 629 d’ Autriche, 629 de fontaine, 629 Crété, M523 Crossbill, 72 Whitewinged, 72 Crossman, E.J., Reviews by, 161,676 Crow, American, 138 Cryptantha shackletteana, 198 spiculifera, 291 Cryptogramma crispa var. sitchensis, 200 stelleri, 198 Cuddy, D., 571 Culaea inconstans, 79 Cumming, E. E. and A. W. Diamond. Songbird Community Composition Versus Forest Rotation Age in Saskatchewan Boreal Mixedwood Forest, 69 Curculio spp., 109 Currant, Northern Black, 465 Cyanocitta cristata, 137 Cygnus buccinator, 368 columbianus, 404 Cynomys ludovicianus, 307 Cyprinus carpio, 30,434 Cypripedium acaule, 601 calceolus ssp. parviflorum, 457 guttatum, 447 parviflorum var. makasin, 447 parviflorum var. pubescens, 575 Cystophora cristata, 607 Cystopteris fragilis, 447 montana, 200,447 Dace, Blacknose, 536 Northern Redbelly, 537 Redside, 76 Striped, 538 Dace, Clinostomus elongatus, in the Lynde and Pringle Creek Watersheds of Lake Ontario, Status of THE CANADIAN FIELD-NATURALIST Vol. 116 - Redside, 76 Dace, Rhinichthys atratulus, in Nova Scotia, Distribution of Blacknose, 536 Dactylis glomerata, 319 Dactylopterus volitans, 134 Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia, Meek’s Halfbeak, Hyporhamphus meeki, and Flying Gurnard, 134 Daigle, C., P. Galois, and Y. Chagnon. Nesting Activities of an Eastern Spiny Softshell Turtle, Apalone spini- fera, 104 Daigle, G., 523 Daisy, Ox-eye, 575 Dalibarda repens, 525 Dandelion, 54,319,575 Common, 473,579 Horned, 473 Northern, 302 Pink, 297 Danthonia intermedia, 200 parryi, 52 sericea, 114 spicata, 575 Darbyshire, S. J. Ephemeral Occurrence of the Mosquito Fern, Azolla caroliniana, at Ottawa, Ontario, 441 Darimont, C. T. and P. C. Paquet. The Gray Wolves, Canis lupus, of British Columbia’s Central and North Coast: Distribution and Conservation Assessment, 416 Darter, 35 Deer, Black-tailed, 416 Mule, 142,488 White-tailed, 123,127,313,315,488,523,648 Deer, Odocoileus virginianus, and Snowshoe Hare, Lepus americanus, in Mixed and Coniferous Forests of Southeastern Québec, Rapid Estimation of Plant Biomass Used as Forage or Cover by White-tailed, 523 Deer, Odocoileus virginianus, in Québec, Testing a Double-Count Aerial Survey Technique for White- Tailed, 488 Delphinium brachycentrum, 200 Dendroica caerulescens, 487 castanea, 487 coronata, 73,407,487 fusca, 487 magnolia, 487 palmarum, 487 striata, 487 tigrina, 487 virens, 72,487 Dennstaedtia punctilobula, 482,525,600 Dentaria lanciniata, 615 Dermochelys coriacea, 553 Deschampsia brevifolia, 200 caespitosa ssp. brevifolia, 212 Descurainia incana, 615 incisa var. incisa, 447,616 pinnata var. brachycarpa, 616 richardsonii, 615 sophia, 616,626 sophioides, 616 Descuria pinnata, 616 DesGranges, J.-L., 551 Desmognathus fuscus, 554 2002 ocrophaeus, 553 Desroches, J.-F. et B. Couture. Extension de |’ aire de distri- bution de la Salamandre 4 quatre doigts, Hemi- dactylium scutatum, dans |’est du Québec, et notes sur habitat, 317 Devil, Glaucous King, 578 King, 575 Diadophis punctatus, 554 Diamond, A. W., 69 Dicranum polysetum, 601 scoparium, 601 Didiuk, A. B., 323 Diervilla lonicera, 525 Diplotaxe 4 feuilles ténues, 626 des murs, 626 fausse-roquett, 626 Diplotaxis erucoides, 626 muralis, 626 tenuifolia, 616,626 Dock, Bering Sea, 234 Curled, 459 Dog, 125,418 Prairie, 307 Dog, Canis familiaris, Activities in Central Yukon, Wolf, Canis lupus, Response to Domestic Sled, 125 Dogfish, Spiny, 120 Dogfish, Squalus acanthias, from the Bay of Fundy, Nova Scotia, A Rare Leucistic Spiny, 120 Dogwood, Red-osier, 468 Douglasia alaskana, 200 arctica, 198 gormanii, 200 Douglasia, Alaskan, 282 Dovekie, 397 Draba adamsii, 618 albertina, 616 alpina, 447,616 arabisans, 616 aurea, 616 bellii, 616 borealis, 447,616 cana, 616 cinerea, 200,616 clivicola, 617 corymbosa, 200,616 crassifolia, 196,616 densifolia, 200,616 exalata, 260 fladnizensis, 196,616 glabella, 617 glabella var. glabella, 617 glabella var. pycnosperma, 617 grandis, 617 groenlandica, 617 hirta, 617 hyperborea, 617 incana, 617 incerta, 200,617 juvenilis, 617 kananaskis, 200,617 lactea, 195,617 lanceolata, 616 lonchocarpa var. lonchocarpa, 617 lonchocarpa var. thompsonii, 199 lonchocarpa vat. vestita, 617 INDEX TO VOLUME 116 693 longipes, 617 macounii, 200,617 macrocarpa, 254 McCallae, 616 murrayi, 617 nemorosa, 617,627 nivalis, 617 norvegica, 617 oblongata, 617 ogilviensis, 618 oligosperma, 200,618 palanderiana, 200 pauciflora, 618 paysonii, 618 porsildii, 200,618 praealta, 618 reptans, 618 ruaxes, 200,618 scotteri, 618 sibirica, 618 stenoloba, 200,447,618 stenopetala, 200 subcapitata, 618 ventosa, 618 ventosa var. ruaxes, 260 verna, 618,627 wahlenbergii, 618 Draba, Alpine, 462 Northern, 462 Dracocephalum parviflorum, 575 Dragonhead, 575 American, 578 Drave des bois, 627 printaniére, 627 Dryopteris sp., 525 carthusiana, 600 cristata, 600 Duck, American Black, 45 Black, 44 Dabbling, 368 Harlequin, 366 Long-tailed, 366,397 Ring-necked, 45 Wood, 45 Dunlin, 397 Duponita fisheri, 84 Eagle, Bald, 603 Ectopistes migratorius, 12 Editor’s Report for Volume 115 (2001), 356 Eedy, W., Reviews by, 167,170,675 Eelgrass, 369 Eider, Common, 22,396 King, 397 Spectacled, 397 Eiders, Somateria mollissima, Nesting in the Digges Sound Region, Nunavut, Status of Common, 22 Elaeagnus angustifolia, 91 commutata, 447 Elaphomyces spp., 102 Elatine triandra, 566 Eleocharis uniglumis, 447 Elk, American, 122,142,183,313,488 Elk, Cervus elaphus, in the French River Delta, Ontario, Anomalies in Incisor Wear of American, 122 694 Elk, Cervus elaphus, in West-Central Alberta, Winter Habitat Selection at Three Spatial Scales by Ameri- can, 183 Elliottia pyroliflorus, 282 Elm, American, 580,593 Elymus calderi, 447 elongatus ssp. ponticus, 446 glaucus, 447 innovatus, 184 macrourus, 447 trachycaulus ssp. glaucus, 447 trachycaulus ssp. novae-angliae, 447 Empetrum nigrum, 196 Empidonax alnorum, 487 flaviventris, 487 Emydoidea blandingii, 554 Enchelyopus cimbrius, 134 Engeman, R. M., 636 Epigaea repens, 525,601 Epilobium angustifolium, 482,525,600 ciliatum ssp. ciliatum, 575 glandulosum, 525 lactiflorum, 200 luteum, 196 Epipactis helleborine, 575 Eptesicus fuscus, 389,645 Eptesicus fuscus, in New Brunswick, Over-wintering and Reproduction by the Big Brown Bat, 645 Equisetum spp., 590 arvense, 196 fluviatile, 196,447 palustre, 447 pratense, 447 scirpoides, 447 sylvaticum, 600 Eremophilia alpestris, 406 Erethizon dorsatum, 649 Erigeron sp., 575 caespitosus, 55,200 compositus, 196 grandiflorus ssp. arcticus, 200 grandiflorus ssp. grandiflorus, 297 humilis, 196 hyssopifolius, 447 lonchophyllus, 447 Eriophorum angustifolium, 81,447 callitrix, 200 vaginatum, 447 viridi-carinatum, 198 Errata The Canadian Field-Naturalist, 354 Eruca vesicaria ssp. sativa, 627 Erucastrum gallicum, 618,627 Erysimum angustatum, 618 arenicola, 618 capitatum, 618 cheiranthoides, 447,618,627 cheiri, 446,627 coarctatum, 618 hieraciifolium, 618,627 inconspicuum, 446,618 pallasii, 200,619 pallasii var. pallasii, 201 repandum, 627 Esox lucius, 434 masquinongy, 433 THE CANADIAN FIELD-NATURALIST —? Aa Ee Ki hii mors, Vol. 116 niger, 539 Esox masquinongy, as a host for the Silver Lamprey, Ichthyomyzon unicuspis, in the Ottawa River, Ontario/Québec, The Muskellunge, 433 Etcheverry, P., 523 Etheostoma spp., 35 caeruleum, 79 nigrum, 79 Eumetopias jubatus, 418 Eupatorium rugosum, 525 Euphrasia mollis, 200 Eurycea bislineata, 554 Eutrema edwardsii, 200,619 Eyebright, Subalpine, 292 a NY sa Fagus grandifolia, 306,485,588 Fairy-slipper, 457 Falco mexicanus, 138 peregrinus, 138,404 rusticolus, 404 sparverius, 487 Falcon, Peregrine, 138,397 Prairie, 138 Fauria crista-galli, 200 Fecske, D. M. and J. A. Jenks. Dispersal by a Male American Marten, Martes americana, 309 Felis concolor, 63,418 Fern, Alaska Parsley, 206 Fragile, 449 Holly, 206 Mosquito, 441 Mountain Bladder, 449 Mountain Fragile, 206 Nahanni Oak, 449 Fern, Azolla caroliniana, at Ottawa, Ontario, Ephemeral Occurrence of the Mosquito, 441 Fescue, Alaska, 212 Baffin, 452 Foothills Rough, 51 Hard, 452 Idaho, 51 Plains Rough, 51 Red, 452 Richardson’s, 214 Rocky Mountain, 214 Rough, 51 Short-leaf, 452 Small-Flower, 214 Tundra, 214,452 Viviparous, 452 Fescue, Festuca campestris, Rangeland to Wildfire, Resili- ence of Foothills Rough, 51 Festuca baffinensis, 447 brachyphylla, 447 brevissima, 200 campestris, 51 hallii, 51 idahoensis, 51 lenensis, 198,447 minutiflora, 197 ovina ssp. alaskensis, 214 richardsonii, 200 rubra, 448 rubra ssp. arctica, 214 rubra ssp. richardsonii, 214 2002 saximontana, 200 trachyphylla, 448 vivipara ssp. glabra, 447 viviparoidea, 81 Festuca campestris, Rangeland to Wildfire, Resilience of Foothills Rough Fescue, 51 Filion, A. A., 122 Finch, Purple, 480 Fir, Balsam, 44,70,476,524,575,593,641 Fish, Buffalo, 30 Fisher, S.J., D.W. Willis, M.M. Olson, and S.C. Krentz. Flathead Chubs, Platygobio gracilis, in the Upper Missouri River: The Biology of a Species at Risk in an Endangered Habitat, 26 Fissurewort, Soft, 264 Flax, Flat-seeded False, 625 Large-seeded False, 625 Small-seeded False, 625 Fleabane, 575 Hyssop-leaf, 472 Large-Flowered, 297 Spear-leaved, 472 Tufted, 296 Flicker, Northern, 480 Flixweed, 626 Flower, Copper, 282 Cuckoo, 626 Flycatcher, Alder, 479 Yellow-bellied, 479 Forbes, G. J., 645 Fox, Arctic, 22 Red, 129,132,418 Fox, Vulpes vulpes, in Ontario: Implications for Disease- spread, Long Distance Movement by a Coyote, Canis latrans, and Red, 129 Foxtail, Meadow, 450 Mountain, 210 Fragaria sp., 525 virginiana, 600 virginiana ssp. virginiana, 575 Fraxinus americana, 524,588 nigra, 598 pennsylvanica, 580 Freedman, B., 475 Frego, K. A., 547 Fritillaria camschatcensis, 196 Frog, Boreal Chorus, 117,553 Green, 554 Mink, 554 Northern Leopard, 552 Pickerel, 554 Western Chorus, 554 Wood, 117,552 Froglog: Newsletter of the Declining Amphibian Popu- lations Task Force, 179,354,512,680 Fuller, W. A. Canada and the “Buffalo”, Bison bison: A Tale of Two Herds, 141 Fulmar, Northern, 397 Fulmarus glacialis, 403 Gabor, T. S., 42 Gadawski, T. R., 42 Gadus morhua, 424 Galium boreale, 55 palustre, 600 INDEX TO VOLUME 116 695 triflorum, 600 Gallinago gallinago, 487 Gallinula chloropus, 444 Galois, P., 104 Gar, Longnose, 434 Garden-rocket, 627 Gasterosteus aculeatus, 537 Gaston, A. J., 22 Gaultheria hispidula, 601 procumbens, 601 Gavia adamsii, 403 immer, 368 pacifica, 368,403 stellata, 403 Gaylussacia baccata, 601 Gehring, T. M., 648 Gentian, Broadpetal, 287 Dane’s Dwarf, 287 Star, 287 Swamp, 287 Gentiana douglasiana, 200 platypetala, 200 Gentianella tenella, 200 Geocaulon lividum, 447 Geothlypis trichas, 487 Geranium bicknellii, 575 Geum aleppicum ssp. strictum, 447 macrophyllum ssp. perincisum, 447 triflorum, 55 Gibson, A. J. F. and R. A. Myers. Meek’s Halfbeak, Hypor- hamphus meeki, and Flying Gurnard, Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia, 134 Gilchrist, H.G., 22 Gilhen, J., 120,632 Gilhen, J. and A. Hebda. Distribution of Blacknose Dace, Rhinichthys atratulus, in Nova Scotia, 536 Gillott, C., Review by, 165 Gingras, B. A., 116 Ginns, J., Review by, 343,506,672 Ginseng, American, 114 Glasswort, Slender, 459 Glaucomys sabrinus, 98 sabrinus griseifrons, 99 Glaucomys sabrinus, in Southeast Alaska, Diets of Northern Flying Squirrels, 98 Glen, W. M., 585 Giyceria pulchella, 200 Goat, Mountain, 313,418 Golden, H. N., 132 Golden-Plover, American, 397 Golden-tuft, 624 Goldeneye, Barrow’s, 366 Common, 45,366 Goldenrod, Canada, 473,575 Early, 575 Gray, 579 Spike-like, 473 Goldenseal, 112 Goldenseal, Hydrastis canadensis, Populations at the Northern Limit of its Range, Recent Trends in Stem Numbers in, 112 Goldenweed, Northern Mock, 472 Goldfish, 434 Goldthread, Trifoliate, 244 696 Goltz, J. P., 645 Gonatus fabricii, 378 Goodwin, C. E., Review by, 164,336,498,658,661 Goodyera repens, 447 Goold, J. C., H. Whitehead, and R. J. Reid. North Atlantic Sperm Whale, Physeter macrocephalus, Strandings on the Coastlines of the British Isles and Eastern Canada, 371 Goose, Brant, 84 Canada, 44,368,397 Greater White-fronted, 397 Ross’, 396 Snow, 397 Gopher, Northern Pocket, 319,322 Gophers, Thomomys talpoides, Evidence of a Second Litter in Northern Pocket, 322 Gophers, Thomomys talpoides, in Alberta Alfalfa Fields, Reproductive Characteristics of Northern Pocket, 319 ; Gosztony Ainley, M., Review by, 508 Grant, T. A., 580 Grapefern, Lanceleaf, 202 Graptemys geographica, 554 Grass spp., 600 Acuminate Panic, 578 Alpine Blue, 453 Annual Blue, 453 Arctic, 81 Arctic Blue, 453 Blue-eyed, 575 Canada Blue, 575 Common Sweet, 452 Glaucous Blue, 453 Glomerate Satin, 578 Indian Rice, 453 Inland Alkalai, 216 Kentucky Blue, 52,453 Large Glume Blue, 216 Northern Blue, 453 Orchard, 319 Polar Alkalai, 216 Poverty Oat, 577 Reed Canary, 442,453 Signal, 81 Slough, 566 Snow, 214 Tall Wheat, 451 Tickle, 575 Timber Wild Oat, 212 Vahl’s Alkalai, 210 Western Rye, 451 Grass, Arctophila fulva, Aquatic Leaves and Regeneration of Last Year’s Straw in the Arctic, 81 Gratiola neglecta, 566 Grebe, Horned, 368 Red-necked, 368 Western, 366 Greenwood (1918-2002), Canadian Orchidologist, A Trib- ute to Edward Warren, 326 Grosbeak, Evening, 72 Pine, 480 Rose-breasted, 70,480 Groundsel, Balsam, 575 Purple-haired, 473 Grouse, Pinnated, 2 THE CANADIAN FIELD-NATURALIST Vol. 116 Ruffed, 2,479 Sage, 1 Sharp-tailed, 1 Grus canadensis, 404 Guillemot, Black, 397 Pigeon, 368 Gull, California, 366 Glaucous, 22,397,605 Glaucous-winged, 366,603 Great Black-backed, 22 Herring, 22,605 Mew, 366 Sabine’s, 397 Thayer’s, 396 Gulls, Larus glaucescens, in the Southern Strait of Georgia, British Columbia, Population Trends of Nesting Glaucous-winged, 603 Gulo gulo, 125,132,418 Gulo gulo, and an American Marten, Martes americana, in Alaska, Predation by Wolves, Canis lupus, on Wol- verines, 132 Gurnard, Flying, 134 Gurnard, Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia, Meek’s Halfbeak, Hyporham- phus meeki, and Flying, 134 Gymnocarpium dryopteris, 600 jessoense ssp. parvulum, 200,447 robertianum, 206 Gyrfalcon, 396 Gyrinophilus porphyriticus, 553 Habenaria, 590 Hackelia deflexa, 200 Haedrich, R.L., 423 Hairgrass, Mountain, 218 Tufted, 212 Halfbeak, Meek’s, 134 Halfbeak, Hyporhamphus meeki, and Flying Gurnard, Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia, Meek’s, 134 Haliaeetus leucocephalus, 603 Halichoerus grypus, 609 Halimolobos mollis, 200,447,619 Halimolobus, Soft, 462 Hall, P., Reviews by, 331,333 Hamatocaulis vernicosus, 81 Hamr, J., F.F. Mallory, I.A. Filion, G.S. Brown and M.A. Jost. Anomalies in Incisor Wear of American Elk, Cervus elaphus, in the French River Delta, Ontario, 122 Haplopappus macleanii, 447 Hare, Snowshoe, 128,523 Hare, Lepus americanus, in Mixed and Coniferous Forests of Southeastern Québec, Rapid Estimation of Plant Biomass Used as Forage or Cover by White-tailed Deer, Odocoileus virginianus, and Snowshoe, 523 Harebell, Arctic, 471 Harper, E. K., 315 Harrier, Northern, 138,582 Hart, J. P. and D. H. Jamieson. Estimating the Weight of Wolves, Canis lupus, from Chest Girth Measure- ments, 313 Hawk, Cooper’s, 137,580 Red-tailed, 137 Rough-legged, 397 2002 Hawk, Accipiter cooperii, Mobbing Black-billed Magpie, Pica hudsonia, Killed by Cooper’s, 137 Hawks, Accipiter cooperii, Successfully Nest at High Densities in the Northern Great Plains, Cooper’s, 580 Hawksbeard, Dwarf, 472 Hawthorn, 92 Hazel, Beaked, 70 Hazlitt, S.L., 603 Heal-all, 575 Common, 578 Heather, Aleutian Mountain, 282 White Mountain, 282 Hebda, A., 536 Hedge-hyssop, Clammy, 566 Helictotrichon hookeri, 55 Helleborine, 575 Helleborine, Common, 578 Hemidactylium scutatum, 317,554 Hemidactylium scutatum, dans |’ est du Québec, et notes sur Vhabitat, Extension de l’aire de distribution de la Salamandre a quatre doigts, 317 Hemlock, 524,598 Eastern, 530 Mountain, 99 Spotted Water, 280 Western, 99 Henry, J.D. and M. Mico. Relative Abundance, Habitat Use, and Breeding Status of Birds in Aulavik National Park, Banks Island, Northwest Territories, 393 Heron, Great Blue, 359,604 Herring, 368 Pacific, 366 Herring, Clupea pallasi, in the Strait of Georgia, British Columbia, Seasonal Distribution of Waterbirds in Relation to Spawning Pacific, 366 Hesperis matronalis, 619,627 Heteropriacanthus cruentatus, 632 Heteropriacanthus cruentatus, Additional Records of Big- eye Fishes (Priacanthidae) from the Atlantic Coast of Nova Scotia, Including the First Record of the Glasseye Snapper, 632 Heterotheca villosa, 55 Hickey, M. B. C., 136 Hickey, M. B. C. Successful Spawning by Chinook Salmon, Oncorhynchus tshawytscha, in the St. Lawrence River at Cornwall, Ontario, 642 Hieracium sp., 525,590 aurantiacum, 601 piloselloides, 575 Hierochloe hirta ssp. arctica, 447 Hipfner, J. M, H. G. Gilchrist, A. J. Gaston and D. K. Cairns. Status of Common Eiders, Somateria mol- lissima, Nesting in the Digges Sound Region, Nunavut, 22 Hippuris montana, 200 Hirth, D. H., J. M. A. Petty, and C. W. Kilpatrick. Black Bear, Ursus americanus, Hair and Apple Trees, Malus pumila, in Northeast North America, 305 Hirundo rustica, 406 Histrionicus histrionicus, 366 Holroyd, G. L. Mobbing Black-billed Magpie, Pica hudso- nia, Killed by Cooper’s Hawk, Accipiter cooperii, 137 INDEX TO VOLUME 116 697 Honesty, 628 Honeysuckle, 91 Hordeum jubatum, 447 Horseradish, 624 Horsetail, Marsh, 449 Meadow, 449 Water, 449 Horungia procumbens, 627 Houston, C.S., Reviews by, 331,349 Houston, C.S. Spread and Disappearance of the Greater Prairie-Chicken, Tympanuchus cupido, on the Canadian Prairies and adjacent areas, | Hudson, R.J., 183 Huettmann, F., Reviews by, 162,169,172,341,347, 504,505,660,673 Hummingbird, Ruby-throated, 479 Hundertmark, K.J., 132 Huperzia lucidula, 600 Hutchinsia procumbens, 627 Hutchinsie couchée, 627 Hyalella azteca, 566 Hydrastis canadensis, 112 Hydrastis canadensis, Populations at the Northern Limit of its Range, Recent Trends in Stem Numbers in Gold- enseal, 112 Ayla versicolor, 554 Hylocomium splendens, 601 Hypericum perforatum, 575 Hyporhamphus meeki, 134 unifasciatus, 134 Hyporhamphus meeki, and Flying Gurnard, Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia, Meek’s Halfbeak, 134 Iberis amara, 627 umbellata, 627 Iberis 4 ombelles, 627 amer, 627 Ichthyomyzon castaneus, 433 unicuspis, 433 Ichthyomyzon unicuspis, in the Ottawa River, Ontario/ Québec, The Muskellunge, Esox masquinongy, as a host for the Silver Lamprey, 433 Ictiobus spp., 30 niger, 434 Ig], L. D. and R. E. Martin. Records of Northern Mocking- bird, Mimus polyglottos, Occurrences in North Dakota During the Twentieth Century, 87 Ilex verticillata, 600 Impatiens sp., 525 capensis, 525,600 noli-tangere, 200 Iridoprocne bicolor, 406 Ironwood, 524,598 Isatis tinctoria, 627 Isoetes echinospora, 200,447 muricata, 202 Jacob’ s-ladder, Northern, 470 Jaeger, Long-tailed, 397 Parasitic, 397 Pomarine, 397 Jamieson, D. H., 313 Jay, Blue, 137 Jenks, J. A., 309 | SS ea ee 698 Jobin, B., D. Rodrigue, and J.-L. DesGranges. Amphibian and Reptile Diversity along the St. Lawrence River, 551 John, R., Reviews by, 160,163,335,338,340,341, 342,499,500,501,502, 511,659 Johnson, G. A. M. and B. Freedman. Breeding Birds in Forestry Plantations and Natural Forest in the Vicinity of Fundy National Park, New Brunswick, 475 Jones, P. F. and R. J. Hudson. Winter Habitat Selection at Three Spatial Scales by American Elk, Cervus ela- phus, in West-Central Alberta, 183 Jost, M. A., 122 Julienne des dames, 627 Junco hyemalis, 407,487 Junco, Dark-eyed, 407,480 Juncus alpinoarticulatus ssp. americanus, 447 balticus var. alaskanus, 447 castaneus ssp. castaneus, 447 castaneus ssp. leucochlamys, 447 filiformis, 200 mertensianus, 200 triglumis ssp. albescens, 447 Jung, T. S., I. D. Thompson, M. B. C. Hickey and R. D. Titman. Apparent Capture Myopathy in Hoary Bats, Lasiurus cinereus: A Cautionary Note, 136 Juniper, 488 Common, 122,575 Creeping, 449 Ground, 449 Juniperus communis, 122,575 communis ssp. depressa, 447 horizontalis, 200,447 osteosperma, 488 virginiana, 92 Kalmia angustifolia, 525,601 latifolia, 530 Kendall, S., 116 Kennedy, C. E., 446 Kestrel, American, 480 Killdeer, 479 Kilpatrick, C.W., 305 Kinglet, Golden-crowned, 72,479 Ruby, 72 Ruby-crowned, 70,479 Kittentails, Northern, 292 Kittiwake, Black-legged, 397 Knotweed, Eastern, 459 Kobresia sibirica, 200 simpliciuscula, 200 Koeleria macrantha, 55 Krentz, S. C., 26 Kuhn, K. M., 389 Kuzyk, G. W. and K. M. Kuzyk. Wolf, Canis lupus, Response to Domestic Sled Dog, Canis familiaris, Activities in Central Yukon, 125 Kuzyk, K. M., 125 l’Erable rouge, 317 Labrador, First Record of an Eastern Coyote, Canis latrans, in, 127 Labrador-tea, 469 Ladies’-tresses, Hooded, 458 Lady’s-slipper, 577 THE CANADIAN FIELD-NATURALIST Vol. 116 Small Yellow, 457 Spotted, 457 Lagopus lagopus, 404 mutus, 404 Lamb’ s-quarters, 459 Lamprey, Chestnut, 433 Sea, 359,436 Silver, 433 Lamprey, [chthyomyzon unicuspis, in the Ottawa River, Ontario/Québec, The Muskellunge, Esox masqui- nongy, as a host for the Silver, 433 Lampropeltis triangulum, 554 Larch, 476 Larix laricina, 476,588 Lark, Horned, 397 Larkspur, Northern, 244 Larus argentatus, 22,605 brachyrhynchos, 366 californicus, 366 glaucescens, 366,603 hyperboreus, 22,406,605 marinus, 22 thayeri, 406 Larus glaucescens, in the Southern Strait of Georgia, British Columbia, Population Trends of Nesting Glaucous-winged Gulls, 603 Lasionycteris noctivagans, 646 Lasiuris borealis, 390 cinereus, 124,136 Lasiurus cinereus: A Cautionary Note, Apparent Capture Myopathy in Hoary Bats, 136 Lasiurus cinereus (Chiroptera: Vespertilionidae), from Prince Edward Island, First record of the Hoary Bat, 124 Lathyrus spp., 322 Lauff, R. F., Review by, 339 Laurel, Mountain, 530 Leafless-Bulrush, Rolland’s, 230 Leatherback, 553 Leatherleaf, 469 Ledum glandulosum, 99 Ledum groenlandicum, 447,525,601 Dee Gr Re 132 Leek, Wild, 114 Lemna, 445 Lemon, M. J. F., 603 Leopardbane, Streambank, 293 Lépidie, 627 a feuilles dissemblables, 628 a feuilles larges, 628 cultivée, 628 de Virginie, 628 densiflore, 628 des champs, 628 perfoliée, 628 Lepidium sp. aucheri, 628 bourgeauanum, 447,619,627 campestre, 619,628 densiflorum, 628 densiflorum var. densiflorum, 619 densiflorum var. elongatum, 619 densiflorum var. macrocarpum, 619 heterophylum, 628 latifolium, 619,628 2002 oxycarpum, 628 perfoliatum, 619,628 ramosissimum, 448,619,628 ruderale, 628 sativum, 628 virginicum, 619,628 Lepisosteus osseus, 434 Lepomis gibbosus, 79 Lepus americanus, 128,523 Lepus americanus, in Mixed and Coniferous Forests of Southeastern Québec, Rapid Estimation of Plant Biomass Used as Forage or Cover by White-tailed Deer, Odocoileus virginianus, and Snowshoe Hare, 523 : Lesquerella arctica, 200,619 arctica ssp. calderi, 447 arenosa, 619 calderi, 619 douglasii, 619 Liatris punctata, 55 Ligusticum scoticum ssp. hultenii, 200 Lilac, 306 Lily, Dwarf Water, 244 Lily-of-the-valley, 575 Wild, 578 Lim, B. K., Review by, 337 Limosella aquatica, 566 Linden, American, 524 Linnaea borealis, 70,196,525,601 borealis ssp. americana, 447 Liochlorophis vernalis, 554 Lion, Mountain, 63 Lithospermum canescens, 549 Lobularia maritima, 628 Locoweed, 54 Field, 467 Huddelson’s, 276 Late Yellow, 276 Loewen, V., 446 Lomatium dissecta, 55 Longspur, Lapland, 397 Lonicera spp., 91 canadensis, 525 Loon, Common Loon, 368 Pacific, 368,397 Red-throated, 397 Yellow-billed, 397 Lophodytes cucullatus, 45 Lota lota, 39 Louse, 649 Lousewort, Capitate, 470 Furbish’s, 114 Langdorf’s, 470 Whorled, 470 Loxia spp., 72 leucoptera, 72 Lucas, Z. N. and D. F. McAlpine. Extralimital Occurrences of Ringed Seals, Phoca hispida, on Sable Island, Nova Scotia, 607 Luetkea pectinata, 196 Lunaria annua, 628 Lupine, 54 Yukon, 276 Lupinus argenteus, 54 kuschei, 198 nootka-tensis, 196 INDEX TO VOLUME 116 699 Lutra canadensis, 417,604 Luxilus cornutus, 79,537 Luzula arctica ssp. arctica, 447 confusa, 447 parviflora ssp. parviflora, 447 rufescens, 447 spicata, 447 Lycopodium annotinum, 600 annotinum var. pungens, 447 clavatum var. monostachyon, 447 obscurum, 600 selago, 447 Lycopus americanus, 525 uniflorus, 600 Lynx canadensis, 126 Lynx, 126 Lysichitum americanum, 99 Lysimachia nummularia, 525 terrestris, 525 Magpie, Black-billed, 137,397 Magpie, Pica hudsonia, Killed by Cooper's Hawk, Accipi- ter cooperii, Mobbing Black-billed, 137 Maher, R., 42 Maianthemum canadense, 525,575,601 stellatum, 198 Maier, T. J. Long-distance Movements by Female White- footed Mice, Peromyscus leucopus, in Extensive Mixed-wood Forest, 108 Makepeace, S., 645 Malcolmia maritima, 628 Malcolmie maritime, 628 Mallard, 45,368 Mallory, F. F.. 122 Mallory, M. L. and A. B. Didiuk. Narwhal, Monodon monoceros, Near Western Baffin Island, Nunavut, Canada, 323 Malus spp., 92 pumila, 305 Malus pumila, in Northeast North America, Black Bear, ‘Ursus americanus, Hair and Apple Trees, 305 Mann, H. and M. V. S. Raju. First Report of the Rare Charophyte Nitella macounii (T. F. Allen) T. PF. Allen in Saskatchewan and Western Canada, 559 Mannagrass, MacKenzie Valley, 214 Maple, Red, 108,476,530,593 Sugar, 476,524,593 Marestail, Mountain, 280 Marine Turtle Newsletter, 178,355,512,680 Marmota monax, 530 Marsh-Marigold, Mountain, 244 Marten, 183 American, 132,309 Marten, Martes americana, Dispersal by a Male American, 309 Marten, Martes americana, in Alaska, Predation by Wolves, Canis lupus, on Wolverines, Gulo gulo, and an American, 132 Martes americana, 132,183,309 Martes americana, Dispersal by a Male American Marten, 309 Martes americana, in Alaska, Predation by Wolves, Canis lupus, on Wolverines, Gulo gulo, and, an Amencan Marten, 132 Martin, R. E., 87 700 Maryland, Bat Night Roost at an Abandoned Mine in Western, 389 Matricaria matricarioides, 448 perforata, 448 Matteuccia struthiopteris, 525 Matthiola bicornis, 628 Matthiola longipetala, 628 Mayweed, Scentless, 472 McAlpine, D. F., 607 McAlpine, D. F., F. Muldoon, and A. I. Wandeler. First record of the Hoary Bat, Lasiurus cinereus (Chiro- ptera: Vespertilionidae), from Prince Edward Island, 124 McAlpine, D. F., F. Muldoon, G. J. Forbes, A. I. Wandeler, S. Makepeace, H. G. Broders, and J. P. Goltz. Over- wintering and Reproduction by the Big Brown Bat, Eptesicus fuscus, in New Brunswick, 645 McCarthy, S. J. J., Review by, 171 McKinstry, M. C. and S. H, Anderson. Survival, Fates, and Success of Transplanted Beavers, Castor canaden- sis, in Wyoming, 60 McNicholl, M. K., Review by, 651 Meadowsweet, Steven’s, 466 Mech, L. D. Breeding Season of Wolves, Canis:lupus, in Relation to Latitude, 139 Mech, L. D. and E. K. Harper. Differential Use of a Wolf, Canis lupus, Pack Territory Edge and Core, 315 Medeola virginiana, 600 Medicago spp., 319,322 lupulina, 575 Medick, Black, 575 Melampyrum lineare, 601 Melandrium affine, 240 macrospermum, 242 Melanitta spp., 368 fusca, 45,366 nigra, 366 perspicillata, 45,366 Melilotus alba, 448 officinalis, 448 Melospiza georgiana, 487 lincolnii, 487 melodia, 487 Menyanthes trifoliata, 447 Merganser, Common, 45,366 Hooded, 45 Red-breasted, 366,396 Mergus merganser, 45,366 serrator, 366,404 Mertensia paniculata var. paniculata, 447 Mice, Peromyscus leucopus, in Extensive Mixed-wood Forest, Long-distance Movements by Female White- footed, 108 Michaud, I. M. G., 408 Mico, M., 393 Micropterus dolomieu, 359,434,539 salmoides, 79 Milfoil, Eurasian Water, 85 Milk-vetch, Alpine, 466 Bodin’s, 466 Elegant, 274 Harrington, 274 Indian, 466 Neglected, 577,640 Nutzotin’s, 274 THE CANADIAN FIELD-NATURALIST —emmmaes iia ram yb | mR |) 9, wp ary tua Vol. 116 Williams’, 276,466 Milkvetch, Astragalus neglectus, in an Eastern Ontario Alvar Woodland, Initial Beneficial Effects of Fire and Bulldozing on Neglected, 640 Mimulus ringens, 525 Mimus polyglottos, 87 Mimus polyglottos, Occurrences in North Dakota During the Twentieth Century, Records of Northern Mockingbird, 87 Minerslettuce, Bostock’s, 234 Mink, 648 Minuartia biflora, 200,447 dawsonensis, 200 michauxii, 575 rossii, 447 stricta, 200 Mitchella repens, 525 Mite, 649 Mitella diphylla, 525 nuda, 525,600 pentandra, 200 Mitrewort, Alpine, 267 Mniotilta varia, 487 Mnium spp., 591 Mockingbird, Northern, 87 Mockingbird, Mimus polyglottos, Occurrences in North Dakota During the Twentieth Century, Records of Northern, 87 Moehringia lateriflora, 196,447 Moneses uniflora, 447 Monnaie-du-pape, 628 Monodon monoceros, 323 Monodon monoceros, Near Western Baffin Island, Nuna- vut, Canada, Narwhal, 323 Monotropa uniflora, 600 Montia bostockii, 200 Moonwort, Mingan, 202 Northeastern, 202 Triangle-Lobe, 202 Moorhens, 444 Moose, 125,127,132,142,417,488 Morone chrysops, 35,434 saxatilis, 434 Morrison, S., Review by, 165 Morton, D., 389 Moss spp., 601 Mountain-trumpet, Narrowleaf, 287 Mouse, 128 Deer, 109 White-footed, 108 Mouse-ear-cress, 624 Moutarde blanche, 630 d’ Inde, 624 des champs, 630 des chiens, 627 des oiseaux, 625 noire, 625 Mud-purslane, 566 Mudpuppy, 554 Mudwort, 566 Mugwort, Aleutian, 472 Michaux’s, 472 Muhlenbergia glomerata, 575 Muldoon, F., 124,645 Mullein, Common, 575 2002 Mulligan, G. A. Chromosome Numbers Determined from Canadian and Alaskan Material of Native and Naturalized Mustards, Brassicaceae (Cruciferae), 611 Mulligan, G. A. Weedy Introduced Mustards (Brassi- caceae) of Canada, 623 Murphy, R. K., 580 Murre, Thick-billed, 397 Murrelet, Marbled, 366 Muskellunge, 433 Muskellunge, Esox masquinongy, as a host for the Silver Lamprey, Ichthyomyzon unicuspis, in the Ottawa River, Ontario/Québec, The, 433 Muskox, 84,400 Mustard, Ball, 629 Black, 625 Blue, 626 Dog, 627 Eastern Tumble, 630 Garlic, 623 Hare’s-ear, 626 Hedge, 630 Indian, 624 Tall Hedge, 630 Tansy, 462 Tower, 624 Treacle, 627 Tumble, 464,630 White, 630 Wild, 630 Wormseed, 462,627 Mustards, Brassicaceae (Cruciferae), Chromosome Num- bers Determined from Canadian and Alaskan Material of Native and Naturalized, 611 Mustards (Brassicaceae) of Canada, Weedy Introduced, 623 Mustela vision, 648 Myagrum perfoliatum, 629 Myagrum perfolie, 629 Myers, R. A., 134 Myotis spp., 136 leibii, 390 lucifugus, 389 Myrica galea, 525 Myriophyllum sibiricum, 85 spicatum, 85 verticillatum, 85,198 Nagoonberry, 466 Najas flexilis, 198 Narwhal, 323 Narwhal, Monodon monoceros, Near Western Baffin Island, Nunavut, Canada, 323 Nasturtium crystallinum, 619,629 microphyllum, 619,629 officinale, 629 National Recovery Plan, 355,681 Navet, 625 Necturus maculosus, 554 Nemopanthus mucronata, 525,601 Nenneman, M. P., R. K. Murphy, and T. A. Grant. Cooper's Hawks, Accipiter cooperii, Successfully Nest at High Densities in the Northern Great Plains, 580 Neotorularia humilis, 614 Nerodia sipedon, 554 INDEX TO VOLUME 116 701 Neslia paniculata, 619,629 Neslie paniculée, 629 Nesodraba grandis, 617 New Brunswick, Breeding Birds in Forestry Plantations and Natural Forest in the Vicinity of Fundy National Park, 475 New Brunswick, Over-wintering and Reproduction by the Big Brown Bat, Eptesicus fuscus, in, 645 Newt, Eastern, 554 Nicholls, J. V., 98 Nighthawk, Common, 479 Night-Heron, Black-crowned, 359 Nitella acuminata, 564 allenii, 564 bastinii, 568 cordobensis, 562 flexilis, 564 hotchkissii, 564 hyalina, 562 macounii, 559 mirabilis, 564 missouriensis, 564 opaca, 564 pseudoflabellata, 563 quadriscutulum, 562 stuartii, 562 terrestris, 562 Nitella, Macoun’s, 559 Nitella macounii (T. F. Allen) T. F. Allen in Saskatchewan and Western Canada, First Report of the Rare Charophyte, 559 North America, Black Bear, Ursus americanus, Hair and Apple Trees, Malus pumila, in Northeast, 305 North Dakota During the Twentieth Century, Records of Northern Mockingbird, Mimus polyglottos, Occur- rences in, 87 Northwest Territories, Relative Abundance, Habitat Use, and Breeding Status of Birds in Aulavik National Park, Banks Island, 393 Norton, M. R., 408 Notophthalmus viridescens, 554 Notropis heterolepis, 538 Nova Scotia, A Rare Leucistic Spiny Dogfish, Squalus acanthias, from the Bay of Fundy, 120 Nova Scotia, Distribution of Blacknose Dace, Rhinichthys atratulus, in, 536 Nova Scotia, Extralimital Occurrences of Ringed Seals, Phoca hispida, on Sable Island, 607 Nova Scotia, Including the First Record of the Glasseye Snapper, Heteropriacanthus cruentatus, Additional Records of Bigeye Fishes (Priacanthidae) from the Atlantic Coast of, 632 Nova Scotia, Meek’s Halfbeak, Hyporhamphus meeki, and Flying Gurnard, Dactylopterus volitans, captured in the Annapolis Basin, 134 Numenius phaeopus, 405 Nunavut, Canada, Narwhal, Monodon monoceros, Near Western Baffin Island, 323 Nunavut, Status of Common Eiders, Somateria mollissima, Nesting in the Digges Sound Region, 22 Nuphar polysepalum, 447 Nuthatch, Red-breasted, 479 Nyctea scandiaca, 406 Nycticorax nycticorax, 359 Nymphaea tetragona ssp. leibergii, 200 702 THE CANADIAN FIELD-NATURALIST Nymphoides peltata, 443 O’ Neil, J., Review by, 333 Oak, Bur, 309,581 Red, 108,530 White, 530 Oak-Fern, Limestone, 206 Oat, Siberian Flase, 453 Oatgrass, Downy, 114 Parry, 52 Siberian, 216 O’Dea, N. R. and R. L. Haedrich. A Review of the Status of the Atlantic Wolffish, Anarhichas lupus, in Canada, 423 Odocoileus hemionus, 142,416,488 virginianus, 123,127,313,315,488,523,648 Odocoileus virginianus, and Snowshoe Hare, Lepus ameri- canus, in Mixed and Coniferous Forests of South- eastern Québec, Rapid Estimation of Plant Biomass Used as Forage or Cover by White-tailed Deer, 523 Odocoileus virginianus, in Québec, Testing a Double- Count Aerial Survey Technique for White-Tailed Deer, 488 Olive, Russian, 91 Olson, M. M., 26 Oncorhynchus mykiss, 79,433 tshawytscha, 642 Oncorhynchus tshawytscha, in the St. Lawrence River at Cornwall, Ontario, Successful Spawning by Chinook Salmon, 642 Onion, Wild, 457 Onobrynchis viciifolia, 448 Onoclea sensibilis, 525,600 Ontario, Abundance and Distribution of Breeding Water- fowl in the Great Clay Belt of Northern, 42 Ontario Alvar Woodland, Initial Beneficial Effects of Fire and Bulldozing on Neglected Milkvetch, Astragalus neglectus, in an Eastern, 640 Ontario, Anomalies in Incisor Wear of American Elk, Cervus elaphus, in the French River Delta, 122 Ontario, Double-crested Cormorants, Phalacrocorax auri- tus, at Inland Lakes North of Lake Huron, 359 Ontario, Ephemeral Occurrence of the Mosquito Fern, Azolla caroliniana, at Ottawa, 441 Ontario: Implications for Disease-spread, Long Distance Movement by a Coyote, Canis latrans, and Red Fox, Vulpes vulpes, in, 129 Ontario Natural Heritage Information Centre Newsletter 6(1) Winter 2002, 178 Ontario/Québec, The Muskellunge, Esox masquinongy, as a host for the Silver Lamprey, /chthyomyzon unicus- pis, in the Ottawa River, 433 Ontario, Status of Redside Dace, Clinostomus elongatus, in the Lynde and Pringle Creek Watersheds of Lake, 76 Ontario, Successful Spawning by Chinook Salmon, Oncorhynchus tshawytscha, in the St. Lawrence River at Cornwall, 642 Opheodrys vernalis, 555 Oporornis philadelphia, 487 Opuntia fragilis, 547 polyacantha, 549 Opuntia fragilis, Bill Dore’s Notes on the Kaladar Cactus, 547 Orchid, Bracted Green, 457 Vol. 116 Northern Bog, 458 Orchis, Round-leaved, 457 Orcinus orca, 324,376 Oreamnos americanus, 313,418 Orthilia secunda, 447 Osmerus mordax, 359 Osmorhiza depauperata, 200 Osmunda cinnamomea, 525,600 claytoniana, 525 regalis, 525 Ostrya virginiana, 524,598 Ottawa Field-Naturalists’ Club Awards for 2001, The, 682 Ottawa Field-Naturalists’ Club Comittee Reports for 2001, The, 515 Ottawa Field-Naturalists’ Club, 8 January 2002, Minutes of the 123rd Annual Business Meeting of The, 514 Ottawa Field-Naturalists’ Club: 7 January 2003, The 124th Annual Business Meeting of The, 179 Ottawa Field-Naturalist’s Club 2003 Council, Call for Nominations: The, 179 Ottawa Field-Naturalists’ Club 2002 Awards, Call for Nominations: The, 179 Otter, River, 417,604 Ovenbird, 70,480 Ovibos moschatus, 84 Owl, Burrowing, 1,307 Great Horned, 138,582 Short-eared, 396 Snowy, 397 Spotted, 183 Owl, Athene cunicularia, Found in Saskatchewan, Large Clutch Size of a Burrowing. 307 Oxalis acetosella, 600 montana, 525 stricta, 525 Oxycoccus microcarpus, 447 Oxytrope, Scamman’s, 278 Oxytropis campestris ssp. jordalii, 200 campestris ssp. varians, 447 huddelsonii, 200 scammaniana, 200 sericea, 54 Paddlefish, 434 Pagophilus groenlandica, 607 Paintbrush, Elegant Indian, 291 Green Indian, 291 Yukon Indian, 292 Panax quinquefolius, 114 Panicum acuminatum var. acuminatum, 575 flexile, 575 Papaver alboroseum, 200 freedmanianum, 249 kluanensis, 249 radicatum ssp. kluanense, 200 walpolei, 200 Paquet, P. C., 416 Parrya arctica, 447,619 nudicaulis, 196,447,620 Parrya, Northern, 464 Partridge, Gray, 16 Parula americana, 487 Parula, Northern, 479 Parus atricapillus, 487 hudsonicus, 487 2002 Passerage des decombres, 628 Passerculus sandwichensis, 407 Passerella iliaca, 407 Pastel des teinturiers, 627 Paszkowski, C. A., G. Scrimgeour, B. A. Gingras, and S. Kendall. A Comparison of Techniques for Assess- ing Amphibian Assemblages on Streams in the Western Boreal Forest, 116 Pea, 322 Pearlwort, Arctic, 240 Pedicularis capitata, 447 furbishiae, 114 langsdorfii ssp. arctica, 447 verticillata, 447 Peeper, Spring, 552 Pennycress, Arctic, 267 Pennyroyal, False, 575 Penstemon gormanii, 200 Pepper-grass, Bourgeau’s, 463,627 Branched, 463 Clasping-Leaved, 628 Common, 628 Field, 628 Forked, 628 Perennial, 628 Poor-man’s, 628 Roadside, 628 Smith’s, 628 Weedy Native, 627 Western, 628 Perdix perdix, 16 Peregrine, 398 Peromyscus leucopus, 108 maniculatus, 109 Peromyscus leucopus, in Extensive Mixed-wood Forest, Long-distance Movements by Female White-footed Mice, 108 Petasites frigidus, 575 frigidus ssp. frigidus, 447 frigidus ssp. palmatus, 447 Petromyzon marinus, 359,436 Petty, J. M. A., 305 Phacelia mollis, 200 Phacelia, Soft, 290 Phalacrocorax auritus, 359,368 pelagicus, 366 pencillatus, 366 Phalacrocorax auritus, at Inland Lakes North of Lake Huron, Ontario, Double-crested Cormorants, 359 Phalaris arundinacea, 442,448 Phalarope, Red, 397 Red-necked, 396 Phalaropus fulicaria, 405 lobatus, 405 Phasianus colchicus, 15 Pheasant, Ring-necked, 15 Phegopteris connectilis, 600 Pheucticus ludovicianus, 487 Phillips, F.R., 127 Phippsia algida, 200 Phleum pratense, 448 Phlox alaskensis, 447 hoodii, 55,200 richardsonii, 200 sibirica ssp. richardsonii, 201 INDEX TO VOLUME 116 703 Phlox, Alaskan, 470 Siberian, 290 Spiny, 287 Phoca hispida, 607 vitulina, 609 Phoca hispida, on Sable Island, Nova Scotia, Extralimital Occurrences of Ringed Seals, 607 Phoxinus eos, 79,537 Phyllodoce aleutica ssp. glanduliflora, 282 glandulijlora, 200 Physaria didymocarpa, 620 Physeter macrocephalus, 371 Physeter macrocephalus, Strandings on the Coastlines of the British Isles and Eastern Canada, North Atlantic Sperm Whale, 371 Pica hudsonia, 137 pica, 406 Pica hudsonia, Killed by Cooper’s Hawk, Accipiter cooperii, Mobbing Black-billed Magpie, 137 Picea abies, 476 glauca, 44,69, 132,184,309,476,524,572,588,641 mariana, 44,132,184,447,476,524,588 rubens, 476,525,588 sitchensis, 99 Pickerel, Chain, 539 Picoides pubescens, 487 villosus, 487 Pigeon, Passenger, 12 Pigmyweed, 464 Pike, Northern, 434 Pimephales promelas, 79 Pine, Australian, 636 Jack, 44,73,476 Lodgepole, 184,450 Pinion, 488 Ponderosa, 309 Red, 598 Shore, 99 White, 108,122,524,530,598,641 Pineappleweed, 472 Pinicola enucleator, 487 Pintail, Northern, 45,368,396 Pinus banksiana, 44,73,476 contorta, 184 contorta var. contorta, 99 contorta ssp. latifolia, 447 edulis, 488 ponderosa, 309 resinosa, 598 strobus, 108,122,524,530,572,598,64 | Pipistrelle, Eastern, 390 Pipistrellus subflavus, 390 Pipit, American, 397 Piranga ludoviciana, 72 Pistia stratiotes, 443 Pixie-Eyes, 286 Placopecten magellanicus, 428 Plantago eriopoda, 200,447 major, 448 Plantain, Common, 470 Saline, 292,470 Platanthera obtusata, 447 Platygobio gracilis, 26 Platygobio gracilis, in the Upper Missouri River: The Biol- ogy of a Species at Risk in an Endangered Habitat, 704 Flathead Chubs, 26 Plectrophenax nivalis, 407 Plethodon cinereus, 554 Pleuropogon sabinei, 81 Pleurozium schreberi, 601 Plover, Black-bellied, 397 Semipalmated, 397 Pluvialis dominica, 405 squatarola, 404 Poa abbreviata ssp. abbreviata, 447 alpina, 447 annua, 448 arctica ssp. arctica compressa, 575 glauca, 196,447 hispidula, 200 macrocalyx, 216 pratensis, 52 pratensis ssp. alpigena, 81 pratensis ssp. pratensis, 448 sandbergii, 55 Podagrostis aequivalis, 200 thurberiana, 210 Podiceps auritus, 368 grisgena, 368 Poecile atricapillus, 138 Point Pelee Natural History News, 355,513,680 Poison-ivy, 578 Polemonium boreale, 447 Pollachius virens, 632 Pollock, 632 Polygala senega, 575 Polygonum buxiforme, 447 coccineum, 27 viviparum, 447 Polyodon spathula, 434 Polystichum lonchitis, 200 commune, 601 Pomatomus saltatrix, 134 Pondweed, 566 Flat-stalk, 450 Flatstem, 210 Fries’, 207 Richardson’s, 450 Sago, 207 Small, 207 White-Stemmed, 207 Yenisei River, 210 Pontederia, 445 Poplar, Balsam, 44,137,184,458,581 Poppy, Arctic, 249 Pale, 248 Walpole’s, 249 Populus spp., 485 balsamifera, 44,137,184,581 balsamifera ssp. balsamifera, 447 tremoloides, 44,69,136,137,184,447,524,572,581 588,641 trichocarpa, 132 Porcupine, 649 Porzana carolina, 408 Potamogeton, 566 berchtoldii, 207 friesii, 201,447 pectinatus, 201 >’ THE CANADIAN FIELD-NATURALIST Vol. 116 praelongus, 201 pusillus var. pusillus, 201 richardsonii, 447 subsibiricus, 198 zosteriformis, 201 Potentilla anserina, 447 argentea, 575 arguta ssp. convallaria, 201 biflora, 201 diversifolia, 195 drummondii, 195 hookeriana ssp. hookeriana var. furcata, 272 litoralis, 201,447 norvegica, 447 palustris, 447 pennsylvanica vat. virgulata, 272 rubricaulis, 201 virgulata, 272 Potvin, F., L. Breton, and L.-P. Rivest. Testing a Double- Count Aerial Survey Technique for White-Tailed Deer, Odocoileus virginianus, in Québec, 488 Prairie-Chicken, Greater, 1 Prairie-Chicken, Tympanuchus cupido, on the Canadian Prairies and adjacent areas, Spread and Disappear- ance of the Greater, 1 Prescott, D. R. C., M. R. Norton and I. M. G. Michaud. Night Surveys of Yellow Rails, Coturnicops noveb- oracensis, and Virginia Rails, Rallus limicola, in Alberta using Call Playbacks, 408 Priacanthus arenatus, 632 Primrose, Bird’s-eye, 469 Gorman’s Dwarf, 282 Greenland, 287 Mackenzie River Dwarf, 282 Primula cuneifolia ssp. saxifragifolia, 201 egaliksensis, 196 mistassinica, 447 Prince Edward Island: A Classification and Ordination Using Multivariate Methods, The Forests of, 585 Prince Edward Island, First record of the Hoary Bat, Lasiurus cinereus (Chiroptera: Vespertilionidae), from, 124 Pristigenys alta, 632 Procyon lotor, 129,582,636 Procyon lotor, Density at an Urban Park, An Extraordinary Raccoon, 636 Pronghorn, 12 Proulx, G. Reproductive Characteristics of Northern Pocket Gophers, Thomomys talpoides, in Alberta Alfalfa Fields, 319 Proulx, G. and P. J. Cole. Evidence of a Second Litter in Northern Pocket Gophers, Thomomys talpoides, 322 Prunella vulgaris, 574 vulgaris ssp. vulgaris, 575 Prunus pensylvanica, 525 serotina, 306,525 Pseudacris crucifer, 552 maculata, 117 triseriata, 554 triseriata maculata, 553 Ptarmigan, Rock, 397 Willow, 397 Pteridium aquilinum, 525,601 aquilinum var. latiusculum, 575 Ptilidium pulcherrimum, 600 2002 Ptilium crista-castrensis, 601 Puccinellia deschampsioides, 201 interior, 201 phryganodes, 81 vahliana, 210 Pussytoes, Pygmy, 471 Rocky Mountain, 293 Showy, 471 Pyare, S., W. P. Smith, J. V. Nicholls, and J. A. Cook. Diets of Northern Flying Squirrels, Glaucomys sabrinus, in Southeast Alaska, 98 Pyrola sp., 525 asarifolia, 447 chlorantha, 447 elliptica, 600 grandiflora, 447 Pyrola, One-flowered, 468 Pyrus spp., 92 Québec, et notes sur I’habitat, Extension de I’ aire de distri- bution de la Salamandre 4 quatre doigts, Hemi- dactylium scutatum, dans |’ est du, 317 Québec, Rapid Estimation of Plant Biomass Used as Forage or Cover by White-tailed Deer, Odocoileus virginianus, and Snowshoe Hare, Lepus ameri- canus, in Mixed and Coniferous Forests of Southeastern, 523 Québec, Testing a Double-Count Aerial Survey Technique for White-Tailed Deer, Odocoileus virginianus, in, 488 Québec, The Muskellunge, Esox masquinongy, as a host for the Silver Lamprey, Jchthyomyzon unicuspis, in the Ottawa River, Ontario/, 433 Quercus spp., 109,530 alba, 530 macrocarpa, 309,581 prinus, 530 rubra, 108 Quillwort, 202 Bristle-like, 448 Raccoon, 129,582,636 Raccoon, Procyon lotor, Density at an Urban Park, An Extraordinary, 636 Radis, 629 sauvage, 629 Radish, 623 Wild, 629 Raifort, 624 Rail, Virginia, 408 Yellow, 408 Rails, Coturnicops noveboracensis, and Virginia Rails, Rallus limicola, in Alberta using Call Playbacks, Night Surveys of Yellow, 408 Rails, Rallus limicola, in Alberta using Call Playbacks, Night Surveys of Yellow Rails, Coturnicops noveb- oracensis, and Virginia, 408 Raja laevis, 428 Raju, M. V. S., 559 Rallus limicola, 408 Rallus limicola, in Alberta using Call Playbacks, Night Surveys of Yellow Rails, Coturnicops novebora- censis, and Virginia Rails, 408 Rana catesbeiana, 554 clamitans, 554 INDEX TO VOLUME 116 705 palustris, 554 pipiens, 552 septentrionalis, 554 sylvatica, 117,552 Rangifer tarandus, 122,127,313,488 Ranunculus acris, 525 cooleyae, 448 cymbalaria, 447 gelidus ssp. grayi, 201 nivalis, 447 pacificus, 201 pedatifidus ssp. affinis, 201 repens, 600 sulphureus var. sulphureus, 248 trichophyllus var. eradicatus, 201 Rape, 625 Bird, 461,623 Raphanus sp., 623 raphanistrum, 620,629 sativus, 629 Rapistre rugueux, 629 vivace, 629 Rapistrum perenne, 629 rugosum, 629 Raspberry, Dwarf, 575 Wild Red, 466 Rattlesnake-plantain, Dwarf, 458 Raven, Common, 394 Recovery: An Endangered Species Newsletter, 177,354, 513,680 Recovery of Nationally Endangered Wildlife 2001-2002 Annual Report, 681 Redcedar, Eastern, 92 Reddoch, A. H., 326 Reddoch, J. M. and A. H. Reddoch. A Tribute to Edward Warren Greenwood (1918-2002), Canadian Orchi- dologist, 326 Redfish, 428 Redpoll, Common, 397 Redstart, American, 397,479 Reedgrass, Marsh, 184 Slimstem, 451 Regulus calendula, 487 satrapa, 487 Reid, R. J., 371 Reindeer, 122 Rempel, R. S., 42 Renaud, C. B. The Muskellunge, Esox masquinongy, as a host for the Silver Lamprey, /chthyomyzon unicus- pis, in the Ottawa River, Ontario/Québec, 433 RENEW: National Recovery Plan Number 21: The Wood Bison, Bison bison athabascae, 178 Response Statements for Extirpated, Endangered and Threatened Species Listed by the Committee on the Status of Endangered Wildlife in Canada (COSE- WIC) in 2001, 681 Rhamnus alnifolius, 525 frangula, 575 Rheum rhaponticum, 448 Rhinichthys atratulus, 79,536 atronasus, 538 cataractae, 79 Rhinichthys atratulus, in Nova Scotia, Distribution of Blacknose Dace, 536 Rhizobium, 441 ee oat ean ee ee Lee ie ee 706 Rhizopogon vinicolor, 101 Rhododendron canadense, 525,601 Rhubarb, 459 Rhus radicans, 525 radicans ssp. negundo, 575 Rhytidiadelphus triquetrus, 600 Ribes sp., 525 hudsonianum, 447 lacustre, 600 Rice, Wild, 82 Rissa tridactyla, 406 Rivest, L.-P., 488 Robin, American, 406,479 Rock-brake, Fragile, 206 Rock-jasmine, 469 Rockcress, 250 Calder’s, 250 Cody’s, 250 Elegant, 461 Holboell’s, 461 Lemmon’s, 254 Nuttall’s, 461 Richardson’s, 462 Smooth Northern, 254 Wall, 624 Rocket, Dame’s, 627 Sea, 625 Turkish, 625 White, 626 Yellow, 624 Rockling, Fourbeard, 134 Rodrigue, D., 551 Roland, C. A., 192 Roquette des jardins, 627 Rorippa amphibia, 620,629 austriaca, 629 barbareifolia, 448,620 calycina, 629 curvipes var. curvipes, 620 curvisiliqua, 620 islandica, 620 palustris, 620 palustris ssp. palustris, 448 sylvestris, 620,630 Rorippa sylvestre, 630 Rosa acicularis, 184,196 acicularis ssp. sayi, 575 arkansana, 55 multiflora, 91 Rosatte, R. C. Long Distance Movement by a Coyote, Canis latrans, and Red Fox, Vulpes vulpes, in Ontario: Implications for Disease-spread, 129 Rose, Little, 270 Multiflora, 91 Prickly, 184,578 Ross, R. K., K. F. Abraham, T. R. Gadawski, R. S. Rempel, T. S. Gabor, and R. Maher. Abundance and Dis- tribution of Breeding Waterfowl in the Great Clay Belt of Northern Ontario, 42 Rouleau, I., M. Crété, G. Daigle, P. Etcheverry, and C. Beaudoin. Rapid Estimation of Plant Biomass Used as Forage or Cover by White-tailed Deer, Odocoi- leus virginianus, and Snowshoe Hare, Lepus ameri- canus, in Mixed and Coniferous Forests of South- eastern Québec, 523 THE CANADIAN FIELD-NATURALIST Vol. 116 Rubus sp., 525 alleganiensis, 530 arcticus ssp. acaulis, 448 canadensis, 600 idaeus, 448 pubescens, 575,600 strigosus, 482 Rumex acetosa ssp. alpestris, 201 beringensis, 198 crispus, 448 graminifolius, 234 Rush, Alpine, 456 Baltic, 456 Chestnut, 456 Merten’s, 232 Thread, 230 Whitish, 456 Rye, Calder’s Wild, 451 Hairy Wild, 184 Thick-spike Wild, 451 Sagebrush, Alaska, 471 Sagesse-des-chirurgiens, 626 Sagina saginoides, 201 Sagittaria, 445 cuneata, 448 Sainfoin, 467 Salamander, Blue-spotted, 554 Dusky, 554 Four-toed, 554 Mountain Dusky, 553 Northern Two-lined, 554 Redback, 554 Spotted, 554 Spring, 553 Salamandre a quatre doigts, 317 Salamandre 4 quatre doigts, Hemidactylium scutatum, dans lest du Québec, et notes sur l’habitat, Extension de l’aire de distribution de la, 317 Salicornia europaea, 448 Salix spp., 137,590 arbusculoides, 448 arctica, 396 barrattiana, 448 commutata, 196 hookeriana, 198 monticola, 459 myrtillifolia, 448 planifolia ssp. pulchra, 448 pseudomonticola, 448 rotundifolia ssp. dodgeana, 201,448 setchelliana, 196 stolonifera, 197 Salmo salar, 537 Salmon, Atlantic, 537 Chinook, 642 Salmon, Oncorhynchus tshawytscha, in the St. Lawrence River at Cornwall, Ontario, Successful Spawning by Chinook, 642 Salvelinus fontinalis, 79,434,537 namaycush, 434 ‘Sambucus racemosa, 482 Sander-Regier, R., Reviews by, 171,509,674 Sanderling, 397 Sandpiper, Baird’s, 397 2002 Buff-breasted, 397 Least, 397 Pectoral, 397 Purple, 396 Semipalmated, 397 Western, 396 White-rumped, 397 Sandrocket, 626 Sandwort, 575 Beautiful, 236 Blunt-leaved, 460 Longstem, 236 Low, 460 Mountain, 460 Rock, 240,578 Sanguisorba officinalis, 448 Sapin baumier, 317 Sapsucker, Yellow-bellied, 479 Sarcoptes scabiei var. canis, 649 Sarsaparilla, Wild, 70,575 Saskatchewan and Western Canada, First Report of the Rare Charophyte Nitella macounii (T. F. Allen) T. F. Allen in, 559 Saskatchewan Boreal Mixedwood Forest, Songbird Com- munity Composition Versus Forest Rotation Age in, Saskatchewan, Large Clutch Size of a Burrowing Owl, Athene cunicularia, Found in, 307 Saskatoon, 465 Saussurea angustifolia ssp. yukonensis, 201 viscida var. yukonensis, 297 Saussurea, Narrowleaved, 297 Savin, Creeping, 207 Saxifraga adscendens ssp. oregonensis, 201 bracteata, 201 foliolosa, 201 lyallii ssp. hultenii, 448 nivalis, 448 oppositifolia, 196 serpyllifolia, 448 sibirica, 267 Saxifrage, Alpine, 465 Leafystem, 267 Red-stemmed, 465 Siberian, 267 Small, 267 Thyme-leaved, 465 Scallop, Giant, 428 Scaup, Greater, 366 Lesser, 45,368 Schellenberg, M.P., Review by, 671 Scheuchzeria, 450 Scheuchzeria palustris ssp. americana, 448 Schoenocrambe linifolia, 620 Schultz, R. N. and P. C. Wilson. Territorial Marking by Lone Male Gray Wolves, Canis lupus, 311 Scircus cyperinus, 482 Scirpus, 445 rollandii, 230 Scolopax minor, 487 Scoter, Black, 366 Surf, 45,366 White-winged, 45,366 Scouring-rush, Dwarf, 449 Scrimgeour, G., 116 INDEX TO VOLUME 116 707 Scutellaria galericulata var. pubescens, 448 parvula, 576 parvula var. parvula, 575 Sea-lovage, Hultén’s, 282 Seal, Grey, 609 Harbour, 609 Harp, 607 Hooded, 607 Ringed, 607 Seals, Phoca hispida, on Sable Island, Nova Scotia, Extralimital Occurrences of Ringed, 607 Sea Lion, Steller, 418 Sea Urchin, Green, 428 Sea Wind: Bulletin of Ocean Voice International 5(1/2) December 2001, 177 Sebastes sp., 428 Seburn, C., Review by, 678 Seburn, D., Reviews by, 497,498,503 Sedge, 54,81,581 Arctic Marsh, 220 Bebb’s, 577 Black-and-White-Scale, 218 Blackish, 226 Blunt, 226 Bristleleaf, 220,575 Bristly-stalked, 224 Buxbaum’s, 218 Chestnut, 575 Circumpolar, 218 Crawford’s, 220 Creeping, 218 Curly, 455 Dunhead, 228 Graceful Mountain, 454 Houghton’s, 577 Inland, 220 Krause’s, 220 Lakeshore, 224 Liddon, 226 Little Green, 228 Low Northern, 454 Meadow, 228 Nard, 224 Northern Show, 455 Parry's, 226 Richardson’ s, 575 Rock, 455 Rockdwelling, 228 Russet, 455 Sheathed, 455 Short-leaved, 454 Siberian Bog, 230 Simple Bog, 230 Single-spike, 455 Sparse-leaved, 455 Spikenard, 454 Spruce Muskeg, 454 Star, 226 Thread-Leaf, 220 Variegated, 228 Water, 454 Weak, 224 Williams’, 230 Wooly-fruit, 224,454 Yellow Bog, 454 708 Seiurus aurocapillus, 487 noveboracensis, 407 Selaginella selaginoides, 448 sibirica, 201 Semotilus atromaculatus, 79,537 Seneca-snakeroot, 575 Senecio sp., 525 atropurpureus, 196 atropurpureus ssp. frigidus, 448 pauperculus, 575 Setophaga ruticilla, 407,487 Shark, Greenland, 607 Shelley, D. P. and T. M. Gehring. Behavioral Modification of Gray Wolves, Canis lupus, Suffering from Sar- coptic Mange: Importance of Sequential Monitor- ing, 648 Shepherd’ s-purse, 461,625 Oval, 627 Shepherdia canadensis, 184,196,448 Shiner, Blacknose, 538 Common, 537 Shoveler, Northern, 45 Sialia sialis, 487 Silene acaulis, 196 acaulis ssp. subacaulescens, 448 involucrata ssp. involucrata, 201 macrosperma, 242 menziesii, 201 menziesii ssp. williamsii, 242 repens, 201 uralensis ssp. uralensis, 201 williamsii, 201 Silverberry, 467 Silverweed, Common, 465 Sinapis alba, 630 arvensis, 620,630 Sinclair, A., 571,640 Sinclair, A. and P. M. Catling. Recent Trends in Stem Numbers in Goldenseal, Hydrastis canadensis, Populations at the Northern Limit of its Range, 112 Siskin, Pine, 73 Sisymbre de Loesel, 630 élevé, 630 officinal, 630 Sisymbrium altissimum, 448,620,630 loesellii, 620,630 officinale, 620,630 orientale, 630 Sisyrinchium sp., 575 Sitta canadensis, 487 Sium suave, 448 Skate, Barndoor, 428 Skilnick, J., 307 Skullcap, 575 Narrow-leaved, 470 Small, 578 Smelowskia borealis, 196,620 calycina, 266 calycina var. americana, 620 calycina var. integrifolia, 620 calycina ssp. integrifolia var. porsildii, 620 calycina ssp. media, 621 calycina var. porsildii, 201,620 media, 621 pyriformis, 621 THE CANADIAN FIELD-NATURALIST Vol. 116 ~ Smelowskia, Northern, 266 Smelt, Rainbow, 359 Smilacina trifolia, 601 Smith, H. T. and R. M. Engeman. An Extraordinary Raccoon, Procyon lotor, Density at an Urban Park, 636 Smith, W. P., 98 Snake, Brown, 554 Common Garter, 552 Milk, 554 Northern Water, 554 Redbelly, 554 Ringneck, 554 Smooth Green, 554 Snapdragon, 575 Dwarf, 577 Snapper, Glasseye, 632 Snapper, Heteropriacanthus cruentatus, Additional Records of Bigeye Fishes (Priacanthidae) from the Atlantic Coast of Nova Scotia, Including the First Record of the Glasseye, 632 Snipe, Common, 479 Snowberry, 575 Soapberry, 467 Sobey, D. G. and W. M. Glen. The Forests of Prince Edward Island: A Classification and Ordination Using Multivariate Methods, 585 Softshell, Spiny, 554 Solidago spp., 482,525,590 canadensis, 575 canadensis var. salebrosa juncea, 575 missouriensis, 55 multiradiata, 196 nemoralis ssp. nemoralis, 575 simplex, 448 Solomon’s Seal, Star-flowered, 232 Somateria fischeri, 404 mollissima, 22,404 mollissima borealis, 22 mollissima sedentaria, 22 mollissima v-nigra, 22 spectabilis, 404 Somateria mollissima, Nesting in the Digges Sound Region, Nunavut, Status of Common Eiders, 22 Somniosus microcephalus, 607 Sora, 408 Sorbus americana, 601 Sorrel, Garden, 234 Sparganium, 566 minimum, 201,448 multipedunculatum, 448 natans, 207 Sparrow, American Tree, 397 Chipping, 73,407 Fox, 397 Harris’ s, 397 Lincoln’s 480 Savannah, 407 Song, 480 Swamp, 480 White-crowned, 397 White-throated, 73,480 Speedwell, Thyme-leaf, 292 Spermophilus spp., 307 2002 Sphagnum, 591 Sphyrapicus varius, 487 Spike-rush, Creeping, 455 Spikemoss, 448 Siberian, 202 Spiraea beauverdiana, 448 latifolia, 482,525 Spiranthes romanzoffiana, 448 Spizella arborea, 407 passerina, 73,407 Springbeauty, Tuberous, 234 Spruce, 132 Black, 44,184,450,476,524,593 Norway, 476 Red, 476,593 Sitka, 99 White, 44,69, 184,309,476,524,593,641 Squalus acanthias, 120 Squalus acanthias, from the Bay of Fundy, Nova Scotia, A Rare Leucistic Spiny Dogfish, 120 Squirrel, Ground, 307 Northern Flying, 98 Prince of Wales Northern Flying, 99 Squirrels, Glaucomys sabrinus, in Southeast Alaska, Diets of Northern Flying, 98 St. John’ s-wort, 575 Common, 578 Staniforth, R.J., W.J. Cody and K.A. Frego. Bill Dore’s Notes on the Kaladar Cactus (Opuntia fragilis), 547 Starwort, Alaska, 243 Long-stalked, 460 Umbrella, 243 Stellaria alaskana, 196 longipes, 448 umbellata, 198 Stercorarius longicaudus, 406 parasiticus, 406 pomarinus, 405 Sterna paradisaea, 406 Sternotherus odoratus, 553 Stevens, M.A. and R.E. Barry. Selection, Size, and Use of Home Range of the Appalachian Cottontail, Sylvilagus obscurus, 529 Stickleback, Threespine, 537 Stickseed, Nodding, 291 Stinkweed, 630 Stipa hymenoides, 448 viridula, 55 Stitchwort, Mountain, 236 Rock, 240 Ross’, 460 Stizostedion vitreum, 35,434 Stock, Evening, 628 Virginia, 628 Storeria dekayi, 554 occipitomaculata, 554 Strawberry, Barren, 579 Scarlet, 578 Strix occidentalis, 183 Strongylocentrotus droebachiensis, 428 Sturgeon, Atlantic, 434 Lake, 434 Subularia aquatica, 201 aquatica var. americana, 621 Sucker, Longnose, 434 INDEX TO VOLUME 116 709 White, 434,537 Sullivan, T. M., R. W. Butler, and W. S. Boyd. Seasonal Distribution of Waterbirds in Relation to Spawning Pacific Herring, Clupea pallasi, in the Strait of Georgia, British Columbia, 366 Sullivan, T. M., S. L. Hazlitt, and M. J. F. Lemon. Popu- lation Trends of Nesting Glaucous-winged Gulls, Larus glaucescens, in the Southern Strait of Georgia, British Columbia, 603 Surfbird, 366 Swallow, Barn, 396 Tree, 397,479 Swan, Trumpeter, 368 Tundra, 396 Sweet-clover, White, 466 Yellow, 466 Sweet-coltsfoot, 575 Palmate-leaf, 578 Sweetroot, Bluntseed, 282 Swertia perennis, 201 Sylvilagus spp., 532 floridanus, 529 obscurus, 529 transitionalis, 529 Sylvilagus obscurus, Selection, Size, and Use of Home Range of the Appalachian Cottontail, 529 Symphoricarpos albus, 575 Synthyris borealis, 196 Syringa vulgaris, 306 Tabouret des champs, 630 Tachycineta bicolor, 487 Tail, Coon’s, 244 Tamarack, 602 Tanager, Western, 72 Taraxacum spp., 319,525 carneocoloratum, 198 ceratophyllum, 448 lyratum, 302 officinale, 54,448,575 phymatocarpum, 201 Taxidia taxus, 307 Taxus canadensis, 600 Tea, Labrador, 99 Teal, Blue-winged, 45 Green-winged, 45 Teesdalia nudicaulis, 630 Teesdalie a tige nue, 630 Tern, Arctic, 397 Thalictrum sp., 525 pubescens, 600 Thamnophis sirtalis, 552 Thellungiella salsuginea, 612 Thermopsis rhombifolia, 54 Thistle, Bull, 575 Canada, 575 Thlaspi arcticum, 196,621 arvense, 448,621,630 Thomomys bottae, 319 talpoides, 319,322 Thomomys talpoides, Evidence of a Second Litter in Northern Pocket Gophers, 322 Thomomys talpoides, in Alberta Alfalfa Fields, Repro- ductive Characteristics of Northern Pocket Gophers, 319 710 Thompson, I. D., 136 Thrift, Sea, 469 Thrush, Hermit, 479 Swainson’s, 70,479 Thuja occidentalis, 122,317,525,572,588,641 plicata, 99 Thuya occidental, 317 Tiarella cordiafolia, 525 Tilia americana, 524 Tillaea aquatica, 464 Timothy, 453 Titman, R. D., 136 Toad, American, 552 Canadian, 117 Western, 117 Todd, L D. and J. Skilnick. Large Clutch Size of a Burrow- ing Owl, Athene cunicularia, Found in Saskatche- wan, 307 Tofieldia coccinea, 448 Tokaryk, T. T., Reviews by, 347,350,509,669,678 Tolypella glomerata, 567 macounii, 564 prolifera, 566 Touch-Me-Not, Western, 278 Tragopogon dubius, 55 Tree, Apple, 305 Treefrog, Gray, 554 Trichodectes canis, 649 Trichophorum pumilum, 198 pumilum var. rollandii, 199 Trichostema brachiatum, 575 Trientalis borealis, 525,600 Trifolium hybridum, 448 hybridum ssp. elegans, 575 pratense, 448,575 repens, 448 Triglochin maritimum, 448 Trillium undulatum, 600 Tringa flavipes, 405 Trisetum sibiricum, 198 sibiricum ssp. litorale, 201 sibiricum ssp. sibiricum, 448 spicatum, 448 Trisetum, Spike, 454 Troglodytes troglodytes, 487 Trout, Brook, 434,537 Lake, 434 Rainbow, 433 Tryngites subruficollis, 405 Tsuga canadensis, 524,530,598 heterophylla, 99 mertensiana, 99 Turdus migratorius, 406,487 Turnstone, Black, 366 Ruddy, 397 Turritis glabra, 613,624 Turtle, Blanding’s, 554 Common Map, 554 Common Musk, 553 Common Snapping, 552 Eastern Spiny Softshell, 104 Painted, 552 Snapping, 106 Spotted, 554 Wood, 553 THE CANADIAN FIELD-NATURALIST Vol. 116 - Turtle, Apalone spinifera, Nesting Activities of an Eastern Spiny Softshell, 104 Twinflower, 70,470 Tympanuchus cupido, | phasianellus, | Tympanuchus cupido, on the Canadian Prairies and adja- cent areas, Spread and Disappearance of the Greater Prairie-Chicken, | Typha, 445 latifolia, 442,566,600 Ulmus americana, 525,580,588 Uria lomvia, 406 Ursus americanus, 63,125,132,305,313,418 arctos, 132,313,418 horribilis, 63 maritimus, 22,313,324 Ursus americanus, Hair and Apple Trees, Malus pumila, in Northeast North America, Black Bear, 305 Usnea spp., 102 Utricularia vulgaris ssp. macrorhiza, 448 Vaccinium spp., 99,525,530 angustifolium, 601 myrtilloides, 601 Vahlodea atropurpurea, 201 Vélar a feuilles d’éperviére, 627 d’ Orient, 626 étale, 627 fausse-giroflée, 627 Verbascum thapsus, 575 Vermivora ruficapilla, 487 Veronica officinalis, 601 serpyllifolia ssp. humifusa, 201 Vetch, Tufted, 575 Vetchling, 54 Viburnum spp., 532 alnifolium, 525 edule, 184,196,448 nudum, 601 Vicia americana, 54 cracca, 575 Viola, sp., 525,590 adunca, 201,575 biflora, 201 blanda, 600 canadensis, 55 cucullata, 600 macloskeyi, 600 nephrophylla, 575 selkerkii, 198 Violet, Great-spurred, 280 Hook-spur, 278,575 Northern Bog, 575 Twoflower, 280 Vireo olivaceus, 487 solitarius, 73,487 Vireo, Blue-headed, 479 Red-eyed, 70,479 Solitary, 73 Vole, 128 Vulpes vulpes, 129,132,418 Vulpes vulpes, in Ontario: Implications for Disease-spread, Long Distance Movement by a Coyote, Canis latrans, and Red Fox, 129 ee 2002 Waiser, B., Reviews by, 346,676 Waldsteinia fragarioides, 575 Wall-rocket, 626 Walleye, 35,434 Wallflower, Arctic False, 464 Common, 462,627 Edwards’ Mock, 264 Pallas’, 262 Shy Wandeler, A. I., 124,645 Warbler, Bay-breasted, 70,479 Black-and-white, 479 Black-throated Blue, 479 Black-throated Green, 70,479 Blackburnian, 70,479 Blackpoll, 479 Canada, 480 Cape May, 72,484 Magnolia, 70,479 Mourning, 480 Nashville, 479 Palm, 479 Tennessee, 70 Wilson’s, 480 Yellow-rumped, 73,407,479 Water-cress, Northern, 629 Southern, 629 Water-Crowfoot, Thread-Leaf, 248 Water-lily, Yellow, 460 Water-parsnip, 468 Water-plantain, Common, 566 Water-starwort, 566 Watermilfoil, Whorlleaf, 280 Waternymph, Wavy, 210 Waterstarwort, Northern, 278 Waterthrush, Northern, 397 Weed, Turnip, 629 Weevil, Acorn, 109 Weseloh, D. V., 359 Whale, Bowhead, 380 Gray, 380 Humpback, 380 Killer, 324,376 Sperm, 371 Whale, Physeter macrocephalus, Strandings on the Coastlines of the British Isles and Eastern Canada, North Atlantic Sperm, 371 Wheatgrass, 52 Bearded, 53 Bluebunch, 58 Northern, 52 Western, 53 Whimbrel, 396 White, K. S., H. N. Golden, K. J. Hundertmark, and G. R. Lee. Predation by Wolves, Canis lupus, on Wolver- ines, Gulo gulo, and an American Marten, Martes americana, in Alaska, 132 Whitefish, Lake, 125,434 Whitehead, H., 371 Whitlow-grass, 627 Alaska, 261,462 Anadyr, 261 Denseleaf, 255 Fewseed, 260 Flattop, 254 INDEX TO VOLUME 116 711 Grayleaf, 254 Longstalk, 258 Palander’s, 260 Porsild’s, 260 Rainier, 260 Snowbed, 255 Yellowstone, 255 Wigeon, American, 45,368,397 Wilhelmsia physodes, 448 Willis, D. W., 26 Willms, W. D., 51 Willow, 137 Barratt’s, 458 Bilberry, 458 Mountain, 459 Northern Bush, 458 Setchell’s, 232 Stoloniferous, 234 Undergreen, 232 Willow-herb, 575 Ciliate, 578 White-Flower, 280 Yellow, 280 Wilsons Pie. 314 Wilsonia canadensis, 487 pusilla, 487 Wintergreen, Arctic, 469 Greenish, 468 One-sided, 468 Pink-flowered, 468 Woad, 627 Wolf, 125,127,132,139,148,311,313,315 Gray, 416,648 Wolf, Canis lupus, Pack Territory Edge and Core, Differ- ential Use of a, 315 Wolf, Canis lupus, Response to Domestic Sled Dog, Canis familiaris, Activities in Central Yukon, 125 Wolffish, Atlantic, 423 Northern, 423 Spotted, 423 Wolffish, Anarhichas lupus, in Canada, A Review of the Status of the Atlantic, 423 Wolverine, 125,132,418 Wolverines, Gulo gulo, and an American Marten, Martes americana, in Alaska, Predation by Wolves, Canis lupus, on, 132 Wolves, Canis lupus, from Chest Girth Measurements, Estimating the Weight of, 313 Wolves, Canis lupus, in Relation to Latitude, Breeding Season of, 139 Wolves, Canis lupus, of British Columbia’s Central and North Coast: Distribution and Conservation Assessment, The Gray, 416 Wolves, Canis lupus, on Wolverines, Gulo gulo, and an American Marten, Martes americana, in Alaska, Predation by, 132 Wolves, Canis lupus, Suffering from Sarcoptic Mange: Importance of Sequential Monitoring, Behavioral Modification of Gray, 648 Wolves, Canis lupus, Territorial Marking by Lone Male Gray, 311 Wood-Pewee, Eastern, 479 Woodcock, American, 479 Woodpecker, Downy, 480 Hairy, 479 2 | 712 Woodrush, Arctic, 456 Confused, 456 Rusty, 456 Small-flowered, 456 Spiked, 456 Wormwood, 471 Northern, 472 Three-Forked, 293 Wren, Winter, 70,479 Wyoming, Survival, Fates, and Success of Transplanted Beavers, Castor canadensis, in, 60 Xema sabini, 406 Yak, 143 Yarrow, Common, 471 Siberian, 471 Index to Book Reviews Botany Brodo, I. M., S. Duran-Sharnoff, and S. Sharnoff. Lichens of North America, 506 Dale, M. R. T. Spatial Pattern Analysis in Plant Ecology, 344 Douglas, G. W., D. Meidinger and J. L. Penny. Rare Native Vascular Plants of British Columbia, Second Edition, 670 Fordin, D. G. Guide to Standard Floras of the World: An Annotated Geographically Arranged Systematic Bibliography of the Principal Floras, Enumerations, Checklists, and Chorological Atlases of Different Areas. Second Edition, 168 Hudler, G. W. Magical Mushrooms, Mischievous Molds, 343 Kershaw, L., J. Gould, D. Johnson, and J. Lancaster. Rare Vascular Plants of Alberta, 344 Lamoureux, G. Flore printaniére, 670 Liebman, M., C. L. Mohler, and C. P. Staver. Ecological Management of Agricultural Weeds, 671 Moore, D., M. M. Nauta, S. E. Evans, and M. Rotheroe. Fungal Conservation: Issues and Solutions, 672 Environment Beazley, K. and R. Boardman. Politics of the Wild: Canada and Endangered Species, 678 Beck, G. G. and B. Littlejohn. Voices for the Watershed: Environmental Issues in the Great Lakes — St. Lawrence Drainage Basin, 676 Chasek, P. S. The Global Environment in the Twenty-First Century: Prospects for International Cooperation, 345 Conrad, N. C. Reading the Entrails: An Alberta Ecohistory, 346 Ford, E. D. Scientific Method for Ecological Research, 171 Foster, D. R. Thoreau's Country: Journey Through a Transformed Landscape, 676 Harrison, P. and F. Pearce. AAAS Atlas of Population and Environment, 170 Hof, J. and M. Bevers. Spatial Optimization for Managed Ecosystems, 346 Lotze, H. and I. Milewski. Two Hundred Years of Eco- THE CANADIAN FIELD-NATURALIST Vol. 116 - Yellow-tuft, 624 Yellowlegs, Lesser, 396 Yellowthroat, Common, 480 Yukon Territory IV, New Records of Vascular Plants in the, 446 Yukon, Wolf, Canis lupus, Response to Domestic Sled Dog, Canis familiaris, Activities in Central, 125 Zizania, 81 palustris, 82 Zonotrichia albicollis, 73,487 leucophrys, 407 querula, 407 Zostera marina, 369 Zygadenus venenosus, 55 system and Food Web Changes in the Quoddy Region, Outer Bay of Fundy, 673 Marchard, P. J. Autumn, A Season of Change, 674 McCloskey, W. Their Fathers' Work: Casting Nets with the World's Fishermen, 169 O'Brien, M. Making Better Environmental Decisions, 171 Poinar, G. Jr., and R. Poinar. The Amber Forest, 347 Reynolds, J. D., G. M. Mace, K. H. Redford, and J. G. Robinson. Conservation of Exploited Species, 677 Terborgh, J., C. van Schaik, L. Davenport, and M. Rao. Making Parks Work: Strategies for Preserving Tropical Nature, 675 Truett, J.C. and S. R. Johnson. The Natural History of an Arctic Oil Field, Development and the Biota, 347 Miscellaneous Boyd, D. R. Northern Wild, Best Contemporary Canadian Nature Writing, 509 Farber, P. L. Finding Order in Nature: The Naturalist Tradi- tion from Linnaeus to E. O. Wilson, 349 Glavin, T. The Last Great Sea: A Voyage through the Human and Natural History of the North Pacific Ocean, 172 Lowman, M. D. Life in the Treetops: Adventures of a Woman in Field Biology, 508 Roskams, S. Excavation, 350 Secord, J. A. Victorian Sensations: The Extraordinary Publication, Reception, and Secret Authorship of Vestiges of the Natural History of Creation, 509 Stein, B. R. On Her Own Terms: Annie Montague Alex- ander and the Rise of Science in the American West, 678 Wilson, D. and D. Elder. Cheltenham in Antarctica: The Life of Edward Wilson, 510 Zoology Andrews, J. S. The Atlas of the Amphibians and Reptiles of Vermont December, 2001, 666 Bezener, A. and K. de Smet. Manitoba Birds, 163 Bibby, C. J., N. J. Burgess, D. A. Hill, and S. H. Mustoe. Bird Census Techniques, 166 Bonaccorso, F. J. Bats of Papua New Guinea, 337 2002 Burton, R. The World of the Hummingbird, 500 Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. W. Kaiser, A. C. Stewart, and M. C. E. McNall. The Birds of British Columbia Volume 4: Passerines [:] Wood-Warblers through Old World Sparrows, 651 Cannings, S.G., L. R. Ramsay, D. F. Fraser, and M. A. Fraker. Rare Amphibians, Reptiles, and Mammals of British Columbia, 655 Clements, J. and N. Shany. A Field Guide to the Birds of Peru, 338 Crother, B.I., J. Boundry, J. A. Campbell, K. de Queiroz, D. R. Frost, R. Highton, J. B. Iverson, P. A. Meylan, T. W. Reeder, M. E. Siedel, J. W. Sites Jr, T. W. Taggart, S.G. Tilley, and D. B. Wake. Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with Comments Regarding Confidence in Our Under- standing, 656 de Juana, E. y J. Varela. Guia de las Aves de Espafia, 342 de la Pena, M. and M. Rumboll. Birds of Southern South America and Antarctica, 341 del Hoyo, J., A. Elliott and J. Sargatal. Handbook of Birds of the World Volume 7 Jacamars to Woodpeckers, 659 Duellman, W. E. Patterns of Distribution of Amphibians: A Global Perspective, 664 Eder, T. and I. Sheldon. Whales and Other Marine Mammals of British Columbia and Alaska, 341 Elphick, C., J. B. Dunning, Jr., and D. A. Sibley. The Sibley Guide to Bird Life and Behavior, 498 Folkens, P. A. Marine Mammals of the Pacific Northwest: A Concise and Comprehensive Waterproof Guide, 162 Folkens, P. A., R. R. Reeves, B. S. Stewart, P. J. Clapham, and J.J. Powell. National Audubon Society Guide to Marine Mammals of the World, 504 Gibbs, D., E. Barnes, and J. Cox. Pigeons and Doves: A Guide to the Pigeons and Doves of the World, 658 Glut, D. F. Dinosaurs, The Encyclopedia. Supplement 2, 669 Guppy, C. S. and J. H. Shepard. Butterflies of British Columbia (including Western Alberta, Southern Yukon, the Alaskan Panhandle, Washington, Northern Oregon, Northern Idaho, Northwestern Montana), 331 Gwynne, D. T. Katydids and Bush-Crickets: Reproductive Behavior and Evolution of the Tettigoniidae, 165 Hall, L. and G. Richards. Flying Foxes: Fruit and Blossom Bats of Australia, 167 Halpern, S. Four Wings and a Prayer, 333 Hess, G. K., R. L. West, M. V. Barnhill, and L. M. Fleming. Birds of Delaware, 339 Howell, S. N. G. Hummingbirds of Norih America: The Photographic Guide, 661 Hulse, A. C., C. J. McCoy, and E. J. Censky. Amphibians and Reptiles of Pennsylvania and the Northeast, 498 INDEX TO VOLUME 116 me Jones, P. H. The Marbled Murrelets of the Caren Range and Middlepoint Bight, 660 Kenward, R. E. A Manual for Wildlife Radio Tagging, 166 Kerr, M. The Canadian Rockies Guide to Wildlife Watch- ing: The Best Places to See and Appreciate Animals in Their Natural Habitat, 333 Krebs, C. J., S. Boutin, and R. Boonstra. Ecosystem Dynamics of the Boreal Forest: The Kluane Project, 331 Kushlan, J. A. and H. Hafner. Heron Conservation, 164 Kuzmin, S. L. The Amphibians of the Former Soviet Union, 665 Lee, J. C. A Field Guide to the Amphibians and Reptiles of the Maya World: The Lowlands of Mexico, North- ern Guatemala, and Belize, 668 Lehman, P. A Birder's Guide to Metropolitan Areas of North America, 499 Lyons, J., P. A. Cochran, and D. Fago. Wisconsin Fishes 2000: Status and Distribution, 161 MacCulloch, R. The Royal Ontario Museum Field Guide to Amphibians and Reptiles of Ontario, 653 Mann, J., R. C. Connor, P. L. Tyack, and H. Whitehead. Cetacean Societies: Field Studies of Dolphins and Whales, 505 Mattison, C. Snake, 667 McGonigal, D. and L. Woodworth. The Complete Encyclo- pedia — Antarctica and the Arctic, 501 Meshaka, W.E. Jr. The Cuban Treefrog in Florida: Life History of a Successful Colonizing Species, 503 Millspaugh, J. J. and J. M. Marzluff. Radio Tracking and Animal Populations, 334 Moriarty, J. J. and A. M. Bauer. State and Provincial Amphibian and Reptile Publications for the United States and Canada, 662 O'Connor, T. The Archaeology of Animal Bones, 669 Orenstein, R, Survivors in Armor: Turtles, Tortoises, and Terrapins, 497 Petranka, J. W. Salamanders of the United States and Canada, 663 Pizzey, G. and F. Knight. The Field Guide to the Birds of Australia, 165 Ridgely, R. and P. Greenfield. The Birds of Ecuador -— Status, Distribution and Taxonomy (Volume 1), Field Guide (Volume II), 335 Rodner, C., M. Lentino and R. Restall. Checklist of the Birds of Northern South America, 340 Russell, A. P. and A. M. Bauer. The Amphibians and Reptiles of Alberta: A Field Guide and Primer of Boreal Herpetology, Second Edition, 654 Shirihai, H., G. Garballo, and A. J. Helbig. Sylvia Warblers, 336 Svensson, L. and P. Grant. The Complete Guide to the Birds of Europe, 502 Swash, A. and R. Still. Birds, Mammals & Reptiles of the Galapagos Islands, 160 Advice for Contributors to The Canadian Field-Naturalist Content The Canadian Field-Naturalist is a medium for the publi- cation of scientific papers by amateur and professional natu- ralists or field-biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All readers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and com- ment items if judged by the Editor to be of interest to read- ers and subscribers, and book reviews. Please correspond with the Book Review Editor concerning suitability of manuscripts for this section. For further information consult: A Publication Policy for the Ottawa Field-Naturalists’ Club, 1983. The Canadian Field-Naturalist 97(2): 231-234. 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Cook, Editor RR 3 North Augusta, Ontario KOG 1RO, Canada 714 TABLE OF CONTENTS (concluded) }00k Reviews OOLOGY: — The Birds of British Columbia Volume 4: Passerines[:] Wood-Warblers through Old World Sparrows — The Royal Ontario Museum Field Guide to Amphibians and Reptiles of Ontario — The Amphibians and Reptiles of Alberta: A Field Guide and Primer of Boreal Herpetology, Second Edition — Rare Amphibians, Reptiles, and Mammals of British Columbia — Scientific and Standard English Names of Amphibians and Reptiles of North America North of Mexico, with Comments Regarding Confidence in Our Understanding — Pigeons and Doves: A Guide to the Pigeons and Doves of the World — Handbook of Birds of the World Volume 7 Jacamars to Woodpeckers — The Marbled Murrelets of the Caren Range and Middlepoint Bight — Hummingbirds of North America: The Photographic Guide — State and Provincial Amphibian and Reptile Publications for the United States and Canada — Salamanders of the United States and Canada — Patterns of Distribution of Amphibians: A Global Perspective — The Amphibians of the Former Soviet Union — The Atlas of the Amphibians and Reptiles of Vermont December, 2001 — Snake — A Field Guide to the Amphibians and Reptiles of the Maya World: The Lowlands of Mexico, Northern Guatemala, and Belize — Dinosaurs, The Encyclopedia. Supplement 2 — The Archaeology of Animal Bones }OTANY: Rare Native Vascular Plants of British Columbia, Second Edition — Flore printaniere — Ecological Management of Agricultural Weeds — Fungal Conservation: Issues and Solutions INVIRONMENT: Two Hundred Years of Ecosystem and Food Web Changes in the Quoddy Region, Outer Bay of Fundy — Autumn, a Season of Change — Making Parks Work: Strategies for Preserving Tropical Nature — Thoreau’s Country: Journey Through a Transformed Landscape — Voices for the Watershed: Environmental Issues in the Great Lakes — St. Lawrence Drainage Basin — Conservation of Exploited Species — Politics of the Wild: Canada and Endangered Species AISCELLANEOUS: On Her Own Terms: Annie Montague Alexander and the Rise of Science in the American West News and Comment ‘roglog: Newsletter of the Declining Amphibian Populations Task Force (53) — Marine Turtle Newsletter (99) — Point Pelee Natural History News 2(3) — Recovery: An Endangered Species Newsletter (22) — National Recovery Plan (23) Long’s Braya (Braya longii Fernald) and Fernald’s Braya (Braya fernaldii Abbe) — Response Statements for Extirpated, Endangered and Threatened Species Listed by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) in 2001 — Recovery of Nationally Endangered Wildlife 2001-2002 Annual Report [he Ottawa Field-Naturalists’ Club Awards April 2001 ndex to Volume 116 Compiled by LESLIE DUROCHER \dvice for Contributors to The Canadian Field-Naturalist Mailing date of the previous issue 116(3) : 15 April 2003 651 670 673 678 680 682 685 714 THE CANADIAN FIELD-NATURALIST Volume 116, Number 4 2002 | Articles : Rapid estimation of plant biomass used as forage or cover by White-tailed Deer, Odocoileus virginianus, and Snowshoe Hare, Lepus americanus, in mixed and coniferous forests of southeastern Quebec ISABELLE ROULEAU, MICHEL CRETE, GAETAN DAIGLE, PIERRE ETHEVERRY, and CHANTAL BEAUCOIN 523 Selection, sizes, site , and use of home range of the Appalachian Cottontail, Sylvilagus obscurus | Monica A. STEVENS and RONALD E. BARRY 529 | Distribution of Blacknose Dace, Rhinichthys atratulus, in Nova Scotia JOHN GILHEN and ANDREW HEBDA 536 | Bill Dore’s notes on the Kaladar Cactus (Opuntia fragilis) : RICHARD J. STANIFORTH, WILLIAM J. Copy, and KATHERINE A. FREGO 547 | Amphibian and reptile diversity along the St. Lawrence River | BENOIT JOBIN, DAVID RODRIGUE, and JEAN-LUC DESGRANGES 551 First report of the rare charophyte Nitella macounii (T. F. Allen) T. F. Allen in Saskatchewan and western Canada ~ HENRY MANN and M. V. S. RAJU 559 Plant community composition and relationships of disturbed and undisturbed alvar woodland PAUL M. CATLING, ADRIANNE SINCLAIR, and DON CUDDY Sf | Cooper’s Hawks, Accipter cooperii, successfully nesting at high densities in the northern Great Plains MELVIN P. NENNEMAN, ROBERT K. MurpPHy, TODD A. GRANT 580 The forests of Prince Edward Island: A classification and ordination using multivariate methods D. G. SoOBEY and W. M. GLEN 585 Population trends of nesting Glaucous-winged Gulls, Larus glaucescens, in the southern Strait of Georgia, British Columbia TERRY M. SULLIVAN, STEPHANIE L. HAZLITT, and Morra J. F. LEMON 603 | Extralimital occurrences of Ringed Seals, Phoca hispida, on Sable Island, Nova Scotia ZOE N. Lucas and DONALD F. MCALPINE 607 | Chromosome numbers determined from Canadian and Alaskan material of native and | naturalized mustards, Brassicaceae (Cruciferae) GERALD A. MULLIGAN 611 Weedy introduced mustards (Brassicaceae) of Canada GERALD A. MULLIGAN 623 Additional records of bigeye fishes (Pricanthidae) from the Atlantic coast of Nova Scotia, including the first record of the Glasseye Snapper, Heteropriacanthus cruentatus BRIAN W. CoAD and JOHN GILHEN 632 An extraordinary Raccoon, Procyon lotor, density at an urban park HENRY T. SMITH and RICHARD M. ENGEMAN 636 Notes Initial beneficial effects of fire and bulldozing on Neglected Milkvetch. Astragalus neglectus, | in an eastern Ontario alvar woodland PAuL M. CATLING and ADRIANNE SINCLAIR 640 | Successful spawning by Chinook Salmon, Oncorhynchus tshawytscha, in the St. Lawrence River at Cornwall, Ontario M. BRIAN C. HICKEY . 642 Over-wintering and reproduction by the Big Brown Bat, Eptesicus fuscus, in New Brunswick DONALD F. MCALPINE, FRANCES MULDOON, GRAHAM J. FORBES, ALEXANDER I. WANDELER, | ScoTT MAKEPEACE, HUGH G. BRODERS, and JAMES P. GOLTZ 645 Behavioral modification of Gray Wolves, Canis lupus, suffering from sarcoptic mange: importance of sequential monitoring DOouGLas P. SHELLEY and THOMAS M. GEHRING 648 SMITHSONIAN INSTITUTION LIBRARIES | 3 9088 01226 6169 Ss SONIAN INSTITUTION LIBRARIES WOAH 01481 6383 fe BL Paha wa ce Venginis aeons Pee a Ow 3m ¥ Ob ae pe ae ne ye WUT BT Ae ge fy? 229 Srey we ette rere Weert Pent tepes TYE er be ae ee ye we athe outa rb ebe arty price pie cree trans ge Cour ge owe: (te ot tes ts ; ty Wks dry wae pe pet ge ied od iow ye ged ios ee wr FW eae we ae te oe = ym gee